CUCURBITACEAE
葫芦科 hu lu ke
1, Huang Luqi (黄璐琦)2, Chen Shukun (陈书坤)3; Charles Jeffrey4

Lu Anmin (路安民 Lu An-ming)

Herbs, annual or perennial, or weak, woody shrubs with watery sap, scandent or prostrate. Roots fibrous or tuberous. Stem often
angular. Leaves alternate, undivided or variously palmately or pedately divided, often cordate; tendrils solitary, lateral, simple orbranched, spirally twisted, rarely absent; stipules absent. Plants monoecious or dioecious; flowers unisexual, very rarely bisexual;
flowers paniculate, racemose, or subumbellate, rarely solitary. Calyx tube (hypanthium) adnate to ovary; tube rotate, campanulate, orsaucer-shaped, usually 5-lobed; segments imbricate. Corolla usually sympetalous, inserted on calyx tube; segments valvate or
involute. Stamens inserted at base or mouth of calyx tube, usually 5 or 3, of which one often 1-celled and other two 2-celled; filaments separate or variously united into a column; anthers separate or coherent into a head; anther cells straight to conduplicate,extrorse; rudimentary ovary often present in male flowers; staminodes often in female flowers. Ovary inferior or nearly completely
so, mostly composed of 3 carpels, 3-locular, rarely 1- or 2- or spuriously 4–6-locular; ovules usually numerous, rarely few or solitary,horizontal, pendulous, or ascending, often immersed in pulp; placentas parietal, fleshy, often confluent at middle of ovary; style terminal, simple or branched at apex or styles free; stigma enlarged or 2-fid. Fruit usually a fleshy berry or corky, indehiscent or rarely
opening by valves or by an operculum, rarely fibrous. Seeds often numerous, rarely few or solitary, horizontal, pendulous, or
ascending, often compressed, rarely winged; endosperm absent; embryo with leaflike cotyledons and short radicle.
About 123 genera and over 800 species: most in tropics and subtropics, very rare in temperate regions; 35 genera (one endemic, nine introduced)
and 151 species (73 endemic, 14 introduced) in China.
Lu An-ming & Zhang Zhi-yun. 1986. Cucurbitaceae. In: Lu An-ming & Chen Shu-kun, eds., Fl. Reipubl. Popularis Sin. 73(1): 84–280.

1a. Corolla segments fimbriate at margin.
2a. Ovules and seeds numerous; calyx tube less than 7 cm .................................................................................... 21. Trichosanthes
2b. Ovules 12, seeds usually 6, each with 1 abortive seed attached to its side; calyx tube 7–10 cm ......................... 20. Hodgsonia
1b. Corolla segments not fimbriate.
3a. Stamens 5, anther cells ovoid and straight.
4a. Leaves palmately compound, rarely simple.
5a. Plants woody, scandent; leaflets almost entire, with 2 glands at base; seeds with membranous
wing at apex ............................................................................................................................................ 4. Neoalsomitra
5b. Plants herbaceous, scandent; leaflets distinctly denticulate, eglandular at base; seeds without
wing or rarely winged.
6a. Fruit indehiscent; seeds horizontal ................................................................................................... 11. Thladiantha
6b. Fruit 3-valved, dehiscent at apex or rarely not dehiscent but then smaller in size; seeds
pendulous.
7a. Corolla segments more than 5 mm; fruit clavate-cylindric, obovoid, or globose, morethan 6-seeded ................................................................................................................................... 1. Hemsleya
7b. Corolla segments less than 3 mm; fruit turbinate or globose, 1–3(–5)-seeded.
8a. Plants monoecious; fruit turbinate, 3-valved, dehiscent at apex ........................................ 2. Gomphogyne
8b. Plants dioecious; fruit globose or campanulate, indehiscent or dehiscent along
ventral suture ....................................................................................................................... 3. Gynostemma
4b. Leaves simple.
9a. Flowers with corolla segments less than 10 mm; fruit dehiscent, operculate or 3-valved.
10a. Leaf blade entire; fruit 6–10 cm, truncate at apex, 3-valved; seeds surrounded by a membranous
wing ........................................................................................................................................................ 5. Zanonia
10b. Leaf blade divided; fruit 1–3.5 cm, operculate; seed not winged or with membranous wingat apex.
11a. Plants monoecious; leaf blade cordate-hastate, eglandular at base; fruit dehiscentnear middle; seeds not winged ............................................................................................ 7. Actinostemma
11b. Plants dioecious; leaf blade with 1 or 2 pairs of glands at base; fruit dehiscentnear apex; seeds with membranous wing at apex ............................................................... 6. Bolbostemma

1 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s

Republic of China.

2 Institute of Chinese Materia Medica, Chinese Academy of Chinese Medicine, 16 Nanxiaojie, Dongzhimen Nei Avenue, Beijing 100700, People’s Republic of China.
3 Kunming Institute of Botany, Chinese Academy of Sciences, 132 Lanhei Road, Heilongtan, Kunming, Yunnan 650204, People’s Republic of China.
4 Apt. 91, Block 5, Pr. Morisa Toreza 102, St. Petersburg 194017, Russia.

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CUCURBITACEAE

9b. Flowers with corolla segments ca. 2 cm, or if smaller then calyx longer than corolla; fruit baccate,indehiscent; seed not winged.
12a. Flowers with calyx longer than corolla; anthers reniform; fruit more than 10 cm ......................... 8. Indofevillea
12b. Flowers with corolla longer than calyx; anthers oblong; fruit less than 10 cm.
13a. Plants with black glandular scales ................................................................................................. 9. Siraitia
13b. Plants without glandular scales.
14a. Tendrils spiraling only above point of branching; anther cells straight ...................... 11. Thladiantha
14b. Tendrils spiraling also below point of branching; anther cells reniform-curved .... 10. Sinobaijiania
3b. Stamens 3 or 1, anther cells horizontal when only 1 stamen, or rarely 5 but then anther cells conduplicate.
15a. Stamens united into a central column and pollen sac fusing into ring; leaf blade pedatipartite ............... 22. Cyclanthera
15b. Stamens 3, or rarely 5 but then anther cells conduplicate; leaves not pedately compound.
16a. Flowers less than 10 mm in diam.
17a. Plants dioecious or rarely flowers bisexual; fruit 3-valved from apex to base whenmature; seeds 1–3, pendulous .................................................................................................. 14. Schizopepon
17b. Plants usually monoecious, rarely dioecious; fruit indehiscent; seeds often horizontal.
18a. Male flowers without pistillode; anther cells conduplicate; male and female flowers
fasciculate in leaf axil ...................................................................................................... 29. Diplocyclos
18b. Male flowers with globose or subulate pistillode; anther cells straight, curved or
conduplicate.
19a. Anther cells incurved or reflexed ................................................................................... 33. Solena
19b. Anther cells straight.
20a. Male flowers in sessile or subsessile fascicles; ovary hispid ................................ 27. Mukia
20b. Male flowers in pedunculate cymes, racemes, or umbels, or solitary; ovary
glabrous or with fine hairs.
21a. Filaments as long as or longer than anthers, ca. 1 mm or more.
22a. Corolla yellow; filaments and staminodes glabrous except
at very base; seeds scrobiculate .................................................. 28. Scopellaria
22b. Corolla white or pale yellow; filaments and staminodes
hairy above base; seeds smooth ...................................................... 25. Zehneria
21b. Filaments shorter than anthers, 0.5–1 mm.
23a. Anther connective not produced beyond thecae; stigmas
glabrous, apically shortly 2-lobed; fruit black when ripe ............. 24. Melothria
23b. Anther connective usually shortly produced beyond thecae;
stigmas papillose, rounded; fruit white or red when ripe ............... 25. Zehneria
16b. Flowers more than 10 mm in diam.
24a. Anther cells straight; calyx tube narrowly funnelform .................................................................. 17. Edgaria
24b. Anther cells conduplicate.
25a. Tendrils absent .................................................................................................................... 13. Ecballium
25b. Tendrils present.
26a. Corolla campanulate.
27a. Leaf blade hirsute, eglandular at base; flowers yellow; fruit large ................. 35. Cucurbita
27b. Leaf blade glabrous, with a few glands at base; flowers white;
fruit ca. 5 cm ...................................................................................................... 30. Coccinia
26b. Corolla rotate or rarely campanulate but then segments almost free.
28a. Calyx tube in male flower elongate, tubular or funnelform.
29a. Flowers white; petiole apex 2-glandular ............................................... 31. Lagenaria
29b. Flowers yellow; leaf blade eglandular at base.
30a. Corolla rotate; leaf blade less than 10 cm ............................ 19. Gymnopetalum
30b. Corolla campanulate; leaf blade over 10 cm.
31a. Leaf blade deeply lobed; seeds many, horizontal ................. 15. Biswarea
31b. Leaf blade undivided; seeds few, pendulous ........... 16. Herpetospermum
28b. Calyx rotate.
32a. Flowers with leaflike bract on pedicel; fruit usually muriculate,often 3-valved when mature ................................................................ 12. Momordica
32b. Flowers without bract on pedicel.
33a. Male inflorescence racemose or subumbellate.
34a. Annual herb; fruit many seeded .................................................. 18. Luffa
34b. Perennial climber; fruit 1-seeded; seed large ......................... 23. SechiumCUCURBITACEAE

333b. Flowers solitary or fascicled.
35a. Calyx segments subleaflike, dentate, reflexed ................... 34. Benincasa
35b. Calyx segments subulate, entire, not reflexed.
36a. Tendrils 2- or 3-fid; connective not produced .............. 32. Citrullus
36b. Tendrils simple; connective produced beyond
anther cells .................................................................... 26. Cucumis

1. HEMSLEYA Cogniaux ex F. B. Forbes & Hemsley, J. Linn. Soc., Bot. 23: 490. 1888.
雪胆属 xue dan shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, perennial, scandent. Roots enlarged, oblate or rarely cylindric. Stem and branches slender or moderately robust. Tendrils
filiform, 2-fid. Leaves pedately (3–)5–9(–11)-foliolate; leaflets narrowly oblong-lanceolate to obovate-lanceolate, membranous or
papery. Plants dioecious. Inflorescence cymose-racemose to paniculate, axillary. Male flowers: calyx tube short; segments ovate toovate-lanceolate, ascending, spreading, rotate, recurved, or reflexed; corolla pale yellow, yellow-green, or orange-red, rotate, discoid,turbinate, bowl-shaped, inverted bowl-shaped, Chinese lanternlike, or umbelliform; segments ovate-lanceolate to broadly ovoid,
herbaceous or subsucculent; stamens 5; filaments exserted. Female flowers: calyx and corolla usually as in male flowers, sometimes
slightly larger; ovary subglobose or conical, smooth or muriculate; styles 3; stigmas 2-lobed. Fruit globose, cylindric, or tubularcylindric, longitudinally 9- or 10-striate or 9- or 10-angular, almost smooth or verrucose. Seeds elliptic or broadly ovate, usually withwoody wings, sometimes wings membranous or absent.
About 27 species: subtropical or tropical Asia; 25 species (21 endemic) in China.

1a. Roots without tuber; seeds with broad membranous wing ....................................................................................... 1. H. graciliflora
1b. Roots with tuber (except possibly for H. heterosperma); seeds without wing or with thin woody wing.
2a. Flower smaller, corolla 3–7 mm in diam.
3a. Corolla bowl-shaped, 5–7 mm in diam.; petals fleshy; fruit conical to cylindric, 2.4–3.5 cm ×
8–10 mm; seeds oblong-elliptic to ovate, with a corky wing ........................................................................... 2. H. delavayi
3b. Corolla laxly patelliform, 3–5 mm in diam.; petals membranous-herbaceous; fruit obconic,
2–4 cm × 12–15 mm; seeds lenticular, ± isodiametric, unwinged .......................................................... 3. H. kunmingensis
2b. Flower larger, corolla 8–15 mm in diam.
4a. Corolla segments spreading, explanate, or ascending, corolla bowl-shaped or rotate.
5a. Seeds strongly biconvex, almost without wing, margin slightly broad, 1–1.3(–2) mm; fruit
conical to terete-elliptic.
6a. Corolla 15–20 cm, yellow-green, segments broadly ovate .............................................................. 6. H. panlongqi
6b. Corolla 10–13 cm, green or pale green, segments strap-shaped; fruit conical or terete-conical.
7a. Corolla segments ascending; flower buds obtuse at apex ...................................................... 4. H. macrocarpa
7b. Corolla segments spreading or reflexed; flower buds acute at apex ............................................. 5. H. mitrata
5b. Seeds oblong, lenticular, or strongly biconvex, margin narrower, less than 1 mm in diam.; fruit
globose, narrowly elliptic, or narrowly conical.
8a. Seeds oblong, with thin corky wing; fruit narrowly elliptic to narrowly conical ............................ 7. H. dipterygia
8b. Seeds biconvex, without wing; fruit globose or conical.
9a. Seeds lenticular; corolla bowl-shaped ............................................................................... 11. H. sphaerocarpa
9b. Seeds ovate to orbicular; corolla rotate.
10a. Fruit globose, pedicel with an articulation ......................................................................... 8. H. amabilis
10b. Fruit conical or cylindric, pedicel without an articulation.
11a. Fruit apex with distinct style; corolla segments lanceolate .............................. 9. H. heterosperma
11b. Fruit apex without distinct style; corolla segments ovate-orbicular ................... 10. H. lijiangensis
4b. Corolla segments reflexed or revolute, corolla umbelliform or ± globose.
12a. Seed margin narrower; corolla red-brown or pale red-brown, base often dark red-brown.
13a. Leaflets 9–11; corolla bowl-shaped, 8–10 mm in diam. ...................................................... 25. H. endecaphylla
13b. Leaflets (3–)5–9.
14a. Corolla Chinese lanternlike.
15a. Fruit elliptic; corolla 10–15 mm in diam. ................................................................ 20. H. chinensis
15b. Fruit ovoid to globose; corolla 20–25 mm in diam. ................................................. 21. H. gigantha
14b. Corolla bowl-shaped (H. dulongjiangensis corolla unknown).
16a. Fruit elliptic, 5–8 cm; seed margin erose .......................................................... 22. H. dolichocarpa
16b. Fruit ovoid to globose, 3–4 cm.

CUCURBITACEAE

417a. Seed margin smooth; fruiting pedicel with an articulation; leaflets
5–7, elliptic-lanceolate ............................................................................ 23. H. macrosperma
17b. Seeds margin erose; fruiting pedicel without an articulation;
leaflets 3–5, rhombic .......................................................................... 24. H. dulongjiangensis
12b. Seed margin smooth; corolla pale yellow or yellow-green.
18a. Fruit globose or subglobose.
19a. Corolla umbelliform, 10–12 mm in diam.; fruiting pedicel with anarticulation ............................................................................................................ 12. H. panacis-scandens
19b. Corolla Chinese lanternlike or bowl-shaped, 6–12 mm in diam.
20a. Corolla Chinese lanternlike; fruit 2–2.5 cm in diam. .............................................. 13. H. emeiensis
20b. Corolla bowl-shaped; fruit 3–3.5 cm in diam. ................................................ 14. H. pengxianensis
18b. Fruit conical or elliptic to claviform-elliptic.
21a. Fruit elliptic or claviform, over 7 cm.
22a. Fruit narrowly elliptic-claviform, distinctly longitudinally costate, 7–8 cm,
pedicel with indistinct articulation ..................................................................... 18. H. chengyihana
22b. Fruit broadly to narrowly elliptic, indistinctly costate, (6–)11–17 cm, pedicel
with distinct articulation .................................................................................... 19. H. zhejiangensis
21b. Fruit conical to elliptic, 3.5–5 cm.
23a. Corolla bowl-shaped, segments carnose, 8–9 mm; fruit elliptic to ovoid ......... 17. H. carnosiflora
23b. Corolla Chinese lanternlike, segments thinner.
24a. Fruit conical, base cuneate; corolla segments 14–17 mm ............................. 15. H. turbinata
24b. Fruit elliptic; base obtuse; corolla segments 10–15 mm ............................. 16. H. ellipsoidea
1. Hemsleya graciliflora (Harms) Cogniaux in Engler,
Pflanzenr. 66(IV. 275. I): 24. 1916.
马铜铃 ma tong ling
Alsomitra graciliflora Harms, Bot. Jahrb. Syst. 29: 602.
1901; Gomphogyne bonii Gagnepain; Hemsleya gracilifloravar. tianmuensis X. J. Xue & H. Yao; H. longgangensis X. X.
Chen & D. R. Liang.
Root fibrous; tuber absent. Leaves pedately 7-foliolate;petiole 1.8–3 cm; leaflets oblong-lanceolate to obovate-lanceolate, 5–10 × 2–3.5 cm; petiolule 4–7 mm. Inflorescence largely
cymose; peduncle 5–20 cm, densely pubescent. Male flowers:
pedicel filiform, 1–2 mm; calyx segments spreading, triangular,
ca. 2 × 1 mm; corolla spreading, yellowish to pale yellowgreen, rotate, 5–6 mm; segments obovate, 3–4 × ca. 2 mm,
thinly membranous; filaments short, ca. 1 mm. Female flowers:
calyx and corolla as in male flowers; ovary narrowly cylindric,base attenuate. Fruit conical, 2.5–3.5 × 1–1.5 cm; fruiting pedicel curved, 5–6 mm. Seeds oblong, compressed, 12–14 × 5–6
mm, with membranous wing, wing 3–4 mm at both ends ofseed, seed body obovate. Fl. Jun–Sep, fr. Aug–Nov. 2n = 28.
Mountain slopes; 500–2400 m. Guangdong, Guangxi, Hubei,
Jiangxi, Sichuan, Zhejiang [Vietnam].
The fruit is used medicinally.

brous, apex acute; corolla orange-yellow to pale yellow, bowlshaped, 5–7 mm in diam.; segments ovate, revolute, 5–6 mm;
filaments short, 0.5–1 mm. Female flowers: slightly larger thanmale flowers; ovary cylindric to conical, 4–5 × 2–2.5 mm. Fruitconical to cylindric, 2.4–3.5 cm × 8–10 mm, base attenuate,
slightly curved; fruiting pedicel 4–7 mm. Seeds dark brown,
narrowly elliptic, compressed, 10–12 × 4–6 mm, with corky
wing; wing 3–4 mm at apex, ca. 1 mm wide on both sides; seedbody broadly elliptic to ovate. Fl. Jul–Sep, fr. Aug–Oct.
● Forests, valleys, mountain slopes; 1600–2200 m. SW Sichuan,
C and W Yunnan.

1a. Leaves subglabrous; corolla segments inmale acute at apex .................................... 2a. var. delavayi
1b. Leaves densely pubescent; corollasegments in male rounded-obtuse
at apex .................................................. 2b. var. yalungensis
2a. Hemsleya delavayi var. delavayi
短柄雪胆(原变种) duan bing xue dan (yuan bian zhong)Gomphogyne delavayi Gagnepain, Bull. Mus. Natl. Hist.
Nat. 24: 373. 1918; Hemsleya brevipetiolata Handel-Mazzetti.
Leaves subglabrous. Corolla segments in male flowers
acute at apex.
● Forests, valleys; 1800–2000 m. SW Sichuan, C and W Yunnan.

2. Hemsleya delavayi (Gagnepain) C. Jeffrey ex C. Y. Wu & Z.
L. Chen, Acta Phytotax. Sin. 23: 134. 1985.

2b. Hemsleya delavayi var. yalungensis (Handel-Mazzetti) C.
Y. Wu & Z. L. Chen, Acta Phytotax. Sin. 23: 134. 1985.

短柄雪胆 duan bing xue dan

雅砻雪胆 ya long xue dan

Tuber oblate to conical. Stem and branchlets slender. Tendrils glabrous, 2-fid. Leaves pedately 5–7-foliolate; petiole 2–5mm; leaflets lanceolate to elliptic-lanceolate, median leaflets 3–
10 cm × 8–15 mm; petiolule 2–3 mm. Plants dioecious. Inflorescence dichotomous, to 30 cm; pedicels filiform, 2–5 mm.
Male flowers: calyx segments ovate-lanceolate, 2–4 mm, gla-

Hemsleya brevipetiolata var. yalungensis Handel-Mazzetti, Symb. Sin. 7: 1059. 1936.
Leaves densely pubescent. Corolla segments in male flowers rounded-obtuse at apex.
● Mountain slopes; 1600–2200 m. SW Sichuan.

CUCURBITACEAE

3. Hemsleya kunmingensis H. T. Li & D. Z. Li, Ann. Bot.
Fenn. 44: 486. 2007.
昆明雪胆 kun ming xue dan
Tuber enlarged, ovoid. Stem slender, tomentose when
young but smooth when older. Tendril glabrous, distally 2-fid.
Leaves pedately 7–9-foliolate; petiole 2–4 cm; leaflets lanceolate to oblong; median leaflets 6–12 × 1–1.5 cm; petiolule 2–4
mm. Plants dioecious. Inflorescence dichasial, 5–30 cm, 10–30flowered; pedicels 5–10 mm. Male flowers: calyx segments
broadly triangular, 3–5 mm, revolute, shortly cuspidate at apex;
corolla yellowish green, laxly patelliform, 3–5 mm in diam.,
herbaceous; segments elliptic, 3–5 mm, evaginate and revoluteinto an anchor shape at apex; filaments 1–2 mm; anthers
whitish. Female flowers: calyx and corolla as in male flowers;
ovary conical; styles 3, 1–2 mm. Fruit many, greenish, longitudinally 10-striate, obconic, 2–4 × 1.2–1.5 cm, almost smooth;
fruiting pedicel 10–15 mm. Seeds many, blackish brown, lenticular, 5–8 mm in diam., 2–3 mm thick, verrucose, unwinged. Fl.
Jun–Sep, fr. Aug–Nov.
● Damp valleys in secondary forests; ca. 2300 m. C Yunnan.

4. Hemsleya macrocarpa (Cogniaux) C. Y. Wu ex C. Jeffrey,
Kew Bull. 36: 739. 1982.
圆锥果雪胆 yuan zhui guo xue dan
Tuber oblate or ovoid. Stem sometimes slightly thickened at nodes. Leaves pedately 7–9-foliolate; petiole 3–7 cm,
sparsely puberulent or subglabrous; leaflets obovate-lanceolate
to elliptic-lanceolate; median leaflets 6–10 × 2.5–4 cm; petiolule 2–6 mm. Inflorescence dichotomously cymose, 4–12 cm;
peduncle and branches slightly flexuous. Male flowers: bud obtuse at apex; calyx segments spreading, ovate-lanceolate, 4–5 ×
2–3 mm; corolla green to pale yellow-green, rotate, 7–15 mm indiam.; segments strap-shaped, 6–7 × 2–3 mm, fleshy, base witha pair of glands; filaments less than 1 mm. Female flowers:
calyx and corolla as in male flowers; ovary conical, 5–7 × 3–4
mm. Fruit campanulate to conical, (3–)4–7 × 3–4 cm, with distinct ribs; fruiting pedicel 8–10 mm. Seeds dark brown, irregularly orbicular, 12–15 × 8–12 mm, 3–4 mm thick, with very
narrow wing. Fl. Jul–Sep, fr. Sep–Nov. 2n = 28.

5Evergreen forests; 1000–2200 m. SW Yunnan [NE India].

4b. Hemsleya macrocarpa var. clavata (C. Y. Wu) D. Z. Li,
Syst. Evol. Hemsleya, 76. 1993.
棒果雪胆 bang guo xue dan
Hemsleya clavata C. Y. Wu, Acta Phytotax. Sin. 23: 124.
1985.
Corolla 7–9 mm in diam. Fruit (4–)6.2(–6.5) cm, (1.5–)
2.5(–3) × as long as broad.
● Broad-leaved forests; 1400–1500 m. SW Yunnan.

4c. Hemsleya macrocarpa var. grandiflora (C. Y. Wu) D. Z.
Li, Syst. Evol. Hemsleya, 76. 1993.
大花雪胆 da hua xue dan
Hemsleya grandiflora C. Y. Wu, Acta Phytotax. Sin. 23:130. 1985.
Corolla 12–15 mm in diam. Fruit (4–)6.2(–6.5) cm, (1.5–)
2.5(–3) × as long as broad.
● Thickets on mountain slopes; ca. 2300 m. NW Yunnan.

5. Hemsleya mitrata C. Y. Wu & Z. L. Chen, Acta Phytotax.
Sin. 23: 128. 1985.
帽果雪胆 mao guo xue dan
Tuber enlarged. Stem and branchlets stout, sparsely puberulent, ultimately glabrous; stem nodes thickened, densely
pubescent. Leaves pedately 5–7-foliolate; petiole 5–8 cm; leaflets lanceolate to elliptic-lanceolate, margin serrate; medianleaflets 5–11 × 2.5–4 cm; petiolule 2–5 mm. Plants dioecious.
Inflorescence peduncle slightly reflexed, 5–11 cm. Male flowers: buds acute at apex; calyx segments lanceolate, ca. 6 × 3
mm; corolla pale yellow-green, rotate, 10–11 mm, fleshy; segments ± strap-shaped, base with 2 distinct glands. Female
flower unknown. Fruit caplike, 4–5 × 3–3.6 cm. Seeds darkbrown, broadly ovoid, 12–13 × 9–10 mm, 3–3.5 mm thick,almost unwinged. Fl. Jul–Oct, fr. Sep–Dec.
● Thickets on mountain slopes; 2400–2700 m. SW Yunnan.

Evergreen forests, thickets on mountain slopes; 1000–2300 m.
NW and SW Yunnan [NE India].

6. Hemsleya panlongqi A. M. Lu & W. J. Chang, Acta Phytotax. Sin. 21: 183. 1983.

1a. Fruit (3–)4.5(–6) cm, (1.3–)1.6(–1.85)
× as long as broad ............................... 4a. var. macrocarpa
1b. Fruit (4–)6.2(–6.5) cm, (1.5–)2.5(–3)× as long as broad.
2a. Corolla 7–9 mm in diam. ................... 4b. var. clavata
2b. Corolla 12–15 mm in diam. ........ 4c. var. grandiflora盘龙七 pan long qi

4a. Hemsleya macrocarpa var. macrocarpa
圆锥果雪胆(原变种) yuan zhui guo xue dan (yuan bianzhong)
Gomphogyne macrocarpa Cogniaux in Engler, Pflanzenr.
66(IV. 275. I): 40. 1916; Hemsleya changningensis C. Y. Wu &Z. L. Chen; H. obconica C. Y. Wu & Z. L. Chen.
Fruit (3–)4.5(–6) cm, (1.3–)1.6(–1.85) × as long as broad.

Tuber oblate-spheroid. Branches slender, sulcate-angular,
densely pubescent at nodes. Tendrils 2-fid. Leaves pedately 7foliolate; petiole 2–4 cm; leaflets elliptic-lanceolate to ovatelanceolate, median leaflets 8–12 × 2–3.5 cm, lateral leaflets
smaller, base cuneate, margin serrate, apex acuminate; petiolule
3–6 mm, densely pubescent. Inflorescence peduncle filiform;
pedicel linear, 12–15 mm. Male flower unknown. Female flowers: bracts very small, subulate; calyx segments 5, ovate-lanceolate, ca. 10 × 3 mm, apex acuminate; corolla pale yellow toyellow-green, rotate, 15–20 mm in diam.; segments spreading,broadly ovate, ca. 12 × 8 mm; ovary ovoid-globose to narrowly
ovoid, 8–11 × 4–6 mm. Fl. Sep.
● Broad-leaved forests; ca. 1800 m. S Sichuan.

CUCURBITACEAE

67. Hemsleya dipterygia Kuang & A. M. Lu, Acta Phytotax.
Sin. 20: 88. 1982 [“dipteriga”].
翼蛇莲 yi she lianHemsleya cissiformis C. Y. Wu.
Tuber enlarged. Stem usually thickened at nodes, densely
pubescent, ultimately glabrous. Tendrils glabrous. Leaves pedately 5–7-foliolate; petiole 2–6 cm; leaflets broadly lanceolate
to rhombic, margin serrate or crenate, median leaflets 7–11 ×
ca. 2.5 cm, lateral leaflets smaller; petiolule 2–4 mm. Inflorescence peduncle and branches filiform. Male flowers: pedicel 5–
10 mm; calyx segments ovate, ca. 7 mm; corolla pale yellow topale yellow-green, rotate to bowl-shaped, 12–13 mm in diam.;
corolla segments spreading, broadly obovate, ca. 8 × 7.5 mm,
apex obtuse; filaments less than 1 mm. Female flower unknown. Fruit oblong-claviform or claviform, 4–5.5 × 0.6–1.7
cm, densely pubescent when immature, glabrous when mature,with indistinct ribs. Seeds dark brown, oblong, ca. 14 × 15 mm,
3–4 mm thick. Fl. Jun–Oct, fr. Aug–Nov.
Broad-leaved forests; 100–1500 m. N Guangxi, S Guizhou, C andS Yunnan [N Vietnam].

8. Hemsleya amabilis Diels, Notes Roy. Bot. Gard. Edinburgh
5: 206. 1912.
曲莲 qu lian
Tuber oblate-spheroid. Stem and branchlets very slender.
Tendrils 8–12 cm. Leaves pedately 5–9-foliolate; petiole 2–4
cm; leaflets lanceolate to narrowly lanceolate, median leaflets
4–5 × 1–1.5 cm. Male flowers dichotomously cymose; peduncle slightly flexuous, 5–15 cm; calyx segments 5, ovate-triangular, 4–5 × ca. 2 mm; corolla pale yellow to pale yellowgreen, rotate, 10–12 mm in diam.; segments spreading, broadly
obovate, 5–6 × 4–5 mm; filaments ca. 2 mm, exserted. Female
flowers in simple dichotomously cymose inflorescence, larger
than male flowers, 11–12(–15) mm in diam.; ovary subglobose,
4–5 mm. Fruit spheroid, 12–20 mm; fruiting pedicel filiform,
2–3 mm. Seeds dark brown, broadly ovate-orbicular, 6–8 × 5–7
mm, 3–4 mm thick, unwinged. Fl. Jun–Oct, fr. Aug–Nov. 2n =28*.
● Forests; 1800–3000 m. SW Sichuan, C and W Yunnan.

9. Hemsleya heterosperma (Wallich) C. Jeffrey, Kew Bull. 36:
739. 1982.
异子雪胆 yi zi xue dan

ers similar to male flowers; peduncle 5–11 cm; ovary conical.
Fruit pale yellow-green, narrowly campanulate or claviform,
12–35 × 6–12 mm, glabrous, with distinct ribs, dehiscent at
apex, 3-seeded in each locule. Seeds small, irregularly ovate,ca. 3 × 2 mm, ca. 1 mm thick. Fl. Sep–Oct, fr. Oct–Nov. 2n =28.
Rain forests, evergreen broad-leaved forests; 100–2300 m. Yunnan [Myanmar, Thailand].

10. Hemsleya lijiangensis A. M. Lu ex C. Y. Wu & Z. L. Chen,
Acta Phytotax. Sin. 23: 129. 1985.
丽江雪胆 li jiang xue dan
Tuber enlarged. Stem and branchlets filiform, sparsely pubescent, ultimately subglabrous. Tendrils apically 2-fid. Leaves
pedately 5–7-foliolate; petiole 2–4 cm; leaflets oblong-lanceolate to narrowly lanceolate; median leaflets 6–12 × 2–2.2 cm;
petiolule 2–4 mm. Male flowers: inflorescence peduncle andbranchlets flexuous, 4–5 cm, densely pubescent; pedicel filiform; calyx segments 5, spreading, ovate-triangular, ca. 4 × 2mm, apex acuminate; corolla pale yellow, rotate, 8–10 mm;
segments spreading, obovate, ca. 5 × 4 mm; filaments ca. 2
mm. Female flowers solitary or few dichotomously cymose inleaf axils; pedicel 5–30 mm; ovary narrowly campanulate orconical, ca. 3 × 2 mm, densely white punctate. Fruit campanulate to conical, 2.2–4 cm × 13–18 mm. Seeds dark brown,
irregularly orbicular, compressed, 4–6 × 4–5 mm, 2–3 mm
thick. Fl. Jul–Sep, fr. Aug–Oct.
● Forests; 2000–3000 m. NW Yunnan.

11. Hemsleya sphaerocarpa Kuang & A. M. Lu, Acta Phytotax. Sin. 20: 87. 1982.
蛇莲 she lian
Tuber oblate-spheroid. Stem and branchlets slender, nodes
densely pubescent. Tendrils 8–24 cm. Leaves pedately 5–9foliolate; petiole 1.6–4 cm; leaflets oblong-lanceolate to broadly
lanceolate; median leaflets 7–16 × 2.5–4 cm. Male flowers in alarge dichotomous cyme, 5–25(–45) cm (varying in different
subspecies); calyx segments ovate-triangular, ca. 4 × 3 mm;
corolla pale yellow-green, bowl-shaped, 7–15 mm in diam.;segments broadly ovate, apex acuminate. Female flowers usually in a short inflorescence; ovary spheroid to ovoid, 3–5 mm.
Fruit spheroid to ovoid, 2–3 cm in diam.; fruiting pedicel 1–9mm, with distinct articulation. Seeds lenticular, 8–9 mm indiam., unwinged. Fl. May–Sep, fr. Jul–Nov. 2n = 28*.

Zanonia heterosperma Wallich, Pl. Asiat. Rar. 2: 29. 1831;
Alsomitra heterosperma (Wallich) M. Roemer; Gomphogyne
heterosperma (Wallich) Kurz.

● Broad-leaved forests; 400–2400 m. E and NE Guangxi, S andSE Guizhou, S Hunan, SE, SW, and WC Yunnan.

Tuber unknown. Stem slender, glabrous. Leaves pedately
3–5-foliolate; petiole 3–4 cm, glabrous; leaflets ovate-lanceolate, base attenuate, apex shortly acuminate; median leaflets 5–7
× ca. 1.5 cm; petiolule 3–6 mm. Plants dioecious. Inflorescencecompound or simple; peduncle slender, slightly flexuous, glabrous. Male flowers: calyx segments subulate, ca. 1 mm, apex
apiculate; corolla pale yellow to pale yellow-green, rotate, 3–
4 mm in diam.; segments ovate-lanceolate, ca. 3 × 1 mm, glabrous, apex acuminate; filaments free, ca. 1 mm. Female flow-

1a. Male inflorescence (14–)20(–44)
cm .................................................. 11b. subsp. megathyrsa
1b. Male inflorescence (5–)9(–15) cm.
2a. Corolla (7–)8(–9) mm indiam.; fruiting pedicel
1–2 mm .............................. 11c. subsp. wenshanensis
2b. Corolla (9–)12(–15) mm indiam.; fruiting pedicel
8–9 mm .............................. 11a. subsp. sphaerocarpa

CUCURBITACEAE

11a. Hemsleya sphaerocarpa subsp. sphaerocarpa
蛇莲(原亚种) she lian (yuan ya zhong)
Inflorescence (5–)9(–15) cm. Corolla (9–)12(–15) mm indiam. Fruiting pedicel 8–9 mm.

7● Evergreen broad-leaved forests; 1700–2400 m. SE Yunnan.

12b. Hemsleya panacis-scandens var. pingbianensis C. Y. Wu& Z. L. Chen, Acta Phytotax. Sin. 23: 137. 1985.
屏边藤三七雪胆 ping bian teng san qi xue dan● Broad-leaved forests; 400–1600 m. E and NE Guangxi, S andSE Guizhou, S Hunan, SE and SW Yunnan.

Corolla deep yellow, umbelliform, ca. 15 mm in diam.;segments entire. Ovary and fruit densely white verrucose.

11b. Hemsleya sphaerocarpa subsp. megathyrsa (C. Y. Wu)
D. Z. Li, Syst. Evol. Hemsleya, 84. 1993.

● Broad-leaved forests; ca. 1700 m. SE Yunnan (Pingbian).

大序雪胆 da xu xue dan

13. Hemsleya emeiensis L. D. Shen & W. J. Chang, Acta Phytotax. Sin. 21: 191. 1983.

Hemsleya megathyrsa C. Y. Wu, Acta Phytotax. Sin. 23:
131. 1985; H. megathyrsa var. major C. Y. Wu & Z. L. Chen.

峨眉雪胆 e mei xue dan

Male inflorescence (14–)20(–44) cm. Corolla 8–12 mm indiam.
● Evergreen broad-leaved forests; 2200–2400 m. SW Yunnan.

11c. Hemsleya sphaerocarpa subsp. wenshanensis (A. M. Lu
ex C. Y. Wu & Z. L. Chen) D. Z. Li, Syst. Evol. Hemsleya, 85.
1993.
文山雪胆 wen shan xue dan
Hemsleya wenshanensis A. M. Lu ex C. Y. Wu & Z. L.
Chen, Acta Phytotax. Sin. 23: 130. 1985.
Inflorescence (5–)9(–15) cm. Corolla (7–)8(–9) mm indiam. Fruiting pedicel 1–2 mm.
● Evergreen forests; 1800–2300 m. SE and WC Yunnan (Jingdong, Wenshan).

12. Hemsleya panacis-scandens C. Y. Wu & Z. L. Chen, Acta
Phytotax. Sin. 23: 135. 1985.
藤三七雪胆 teng san qi xue dan
Tuber oblate-spheroid. Stem and branchlets robust, stem
nodes stout, sparsely pubescent. Tendrils 8–15 cm. Leaves pedately 7–9-foliolate; petiole 5–8 cm; leaflets obovate-lanceolate; median leaflets 6–10 × 2–3 cm; petiolule 4–8 mm. Male
flowers dichotomously cymose or paniculate; peduncle usually
filiform, 3–7 cm; calyx segments ovate-lanceolate, ca. 9 × 4mm, apex reflexed; corolla pale yellow to yellow-green, umbelliform, 10–12 mm in diam.; segments obovate to ovate-oblong, fleshy, base thickened, apex obtuse; filaments ca. 1 mm.
Female flowers: corolla 12–15 mm in diam., ovary subglobose,
5–8 × 4–7 mm. Fruit globose, 25–45 mm in diam., fruiting
pedicel 4–8 mm, with articulation. Seeds dark brown, ovoidspheroid, ca. 15 × 12–13 mm, almost unwinged, margin ca. 3
mm wide. Fl. Aug–Oct, fr. Oct–Dec. 2n = 42*.
● Evergreen broad-leaved forests; 1700–2400 m. SE Yunnan.

1a. Corolla pale yellow, segments
involute (not reflexed), marginsinuate ..................................... 12a. var. panacis-scandens
1b. Corolla deep yellow,
umbelliform, margin entire ............ 12b. var. pingbianensis
12a. Hemsleya panacis-scandens var. panacis-scandens
藤三七雪胆(原变种) teng san qi xue dan (yuan bian zhong)Corolla pale yellow, spheroid, 10–12 mm in diam.; segments sinuate at margin. Fruit smooth, 10-angular.

Tuber enlarged, pale brown-yellow. Branchlets slender,
sparsely pubescent when young. Leaves pedately 7–9-foliolate;petiole 4–6 cm; leaflets oblong-lanceolate to ovate-lanceolate;
median leaflets 9–14 × 2–4.5 cm. Male flowers dichotomously
cymose, peduncle 3–8 cm; calyx segments 5, lanceolate, ca. 4 ×
2 mm, apex reflexed; corolla yellow-green, Chinese lanternlike,ca. 8 mm in diam.; segments 5, obovate-lanceolate, ca. 8 × 5mm; filaments ca. 1 mm. Female inflorescence simple; ovary
subglobose, 3–6 mm. Fruit globose, 20–25 mm in diam. Seeds
ovoid, ca. 10 × 8 mm, ca. 3 mm thick, unwinged. Fl. Jul–Sep,
fr. Sep–Nov. 2n = 28*.
● Broad-leaved forests; 1800–2000 m. Sichuan (Emei Shan).

14. Hemsleya pengxianensis W. J. Chang, Acta Phytotax. Sin.
17(4): 97. 1979.
彭县雪胆 peng xian xue dan
Tuber oblate-spheroid. Stem and branchlets slender.
Leaves pedately 5–7-foliolate; petiole 4.5–7 cm, glabrous or
pubescent; leaflets obovate-lanceolate to narrowly lanceolate,
median leaflets 8–17 × 2–4 cm; petiolule 3–5 mm. Male flowers dichotomously cymose; peduncle 4–10 cm, slightly flexuous; calyx segments lanceolate, ca. 4 × 2–3 mm; corolla yellow-green, bowl-shaped or Chinese lanternlike, 6–12 mm indiam.; segments ovate, ca. 6 × 4 mm, apex reflexed. Female
inflorescence reduced; ovary ovoid or subglobose, ca. 3 mm indiam. Fruit ovoid or ellipsoid, 3–4 × 3–3.5 cm. Seeds almost
circular, compressed, 10–13 × 10–13 mm, almost unwinged,
margin 2–3 mm wide. Fl. Jun–Sep, fr. Aug–Nov.
● Broad-leaved forests, forest margins; 700–2100 m. Chongqing,
Sichuan.

1a. Fruit elliptic; corolla Chineselanternlike, 10–12 mm in diam. .......... 14c. var. gulinensis
1b. Fruit ovate to ovoid or subglobose;
corolla bowl-shaped, ca. 8 mm in diam.
2a. Ovary ovoid; petiole, rachis,and corolla pubescent ............... 14b. var. junlianensis
2b. Ovary subglobose; petiole
and corolla glabrous .............. 14a. var. pengxianensis
14a. Hemsleya pengxianensis var. pengxianensis
彭县雪胆(原变种) peng xian xue dan (yuan bian zhong)Hemsleya pengxianensis var. jinfushanensis L. D. Shen &W. J. Chang; H. pengxianensis var. polycarpa L. D. Shen & W.
J. Chang.

CUCURBITACEAE

8

Petiole glabrous. Corolla bowl-shaped, ca. 8 mm in diam.,
glabrous.
● Broad-leaved forests; 700–2100 m. Chongqing, C and SW Sichuan.

leathery. Seeds almost circular, compressed, ca. 15 mm indiam., ca. 3 mm thick, unwinged, margin 3–4 mm wide. Fl.
Jul–Sep, fr. Sep–Nov.
● Broad-leaved forests; ca. 2000 m. Sichuan (Emei Shan).

14b. Hemsleya pengxianensis var. junlianensis L. D. Shen &W. J. Chang, Acta Phytotax. Sin. 21: 188. 1983.

17. Hemsleya carnosiflora C. Y. Wu & Z. L. Chen, Acta
Phytotax. Sin. 23: 133. 1985.

筠连雪胆 jun lian xue dan

肉花雪胆 rou hua xue dan

Petiole, peduncle, and corolla pubescent. Corolla bowlshaped, ca. 8 mm in diam. Ovary ovoid. Fruit ovate to ovoid.

Tuber oblate-spheroid. Stem and branchlets rather robust,thickened at nodes, densely pubescent. Tendrils 7–16 cm.
Leaves pedately 7–9-foliolate; petiole 5–8 cm; leaflets obovatelanceolate, margin dentate; median leaflets 8–12 × 3–4 cm;
petiolule 4–8 mm. Male flowers dichotomously cymose or
paniculate; peduncle indistinctly flexuous, 3–7 cm; calyx segments ovate-lanceolate, ca. 7 × 4 mm, apex reflexed; corolla
pale yellow-green, bowl-shaped, 8–10 mm. Female inflorescence simple; peduncle 2.5–5 cm; corolla larger than male, 11–
12 mm in diam.; ovary ellipsoid, ca. 10 cm × 6–7 mm. Fruitellipsoid to ovoid, 4–5 × 2.5–3 cm; fruiting pedicel 9–12 mm.
Seeds obovate-elliptic to almost circular, compressed, 13–16 ×
12–14 mm, 3–4 mm thick, almost unwinged, margin ca. 4 mm
wide. Fl. Jun–Oct, fr. Aug–Nov. 2n = 42*.

● Forest margins; ca. 1500 m. SW Sichuan.

14c. Hemsleya pengxianensis var. gulinensis L. D. Shen & W.
J. Chang, Acta Phytotax. Sin. 21: 188. 1983.
古蔺雪胆 gu lin xue dan
Hemsleya gulinensis L. D. Shen & W. J. Chang.
Corolla Chinese lanternlike, 10–12 mm in diam. Fruitelliptic.
● Broad-leaved forests. Sichuan.
Hemsleya pengxianensis var. gulinensis is not a new combination
based on H. gulinensis (L. D. Shen & W. J. Chang, Acta Acad. Med.
Sichuan. 11(2): 103. 1980), although both names have the same type.

15. Hemsleya turbinata C. Y. Wu, Acta Phytotax. Sin. 23: 140.
1985.

● Forests; 1800–1900 m. SE Yunnan.

18. Hemsleya chengyihana D. Z. Li, Syst. Evol. Hemsleya, 91.
1993.

陀罗果雪胆 tuo luo guo xue dan

征镒雪胆 zheng yi xue dan

Tuber oblate-spheroid. Stem and branchlets robust. Tendrils 12–20 cm. Leaves 5–7-foliolate; petiole 3–5 cm; leaflets
obovate-lanceolate to elliptic-lanceolate, margin dentate; median leaflets 6–12 cm; petiolule 2–5 mm. Male flowers unknown.
Female flowers 2 to more in a dichotomous cyme; peduncle 3–
10 cm; calyx segments lanceolate, 11–13 × ca. 4 mm, apex
reflexed; corolla pale yellow-green, Chinese lanternlike, 10–12mm; segments oblong-lanceolate, 14–17 × 6–8 mm, base with 2
glands, revolute, acute at apex; ovary conical, ca. 9 × 3–4 mm.
Fruit conical, 3.5–5 × 2–2.5 cm, base attenuate; fruiting pedicel
only ca. 3 mm. Seeds dark brown, broadly ovate, compressed,
9–11 × 8–10 mm, 2–3 mm thick, unwinged, margin ca. 2 mm
wide. Fl. Jul–Sep, fr. Sep–Nov.

Hemsleya macrocarpa C. Y. Wu, Acta Phytotax. Sin. 23:135. 1985, not (Cogniaux) C. Y. Wu ex C. Jeffrey, Kew Bull.
36: 739. 1982.

● Evergreen broad-leaved forests; 1600–2400 m. SW Yunnan.

16. Hemsleya ellipsoidea L. D. Shen & W. J. Chang, Acta Phytotax. Sin. 21: 185. 1983.
椭圆果雪胆 tuo yuan guo xue dan
Tuber enlarged. Stem and branchlets slender, sulcate.
Leaves pedately 3–5-foliolate; petiole 2–5 cm; leaflets ovatelanceolate; median leaflets 9–15 × 4–5 cm; petiolule 3–5 cm.
Male flowers sparsely dichotomously cymose; peduncle 10–12cm; calyx segments lanceolate, ca. 10 × 3 mm, apex reflexed;
corolla pale yellow-green, lantern-shaped, 9–11 mm; segments
broadly ovate-lanceolate, 10–15 × 6–10 mm, apex reflexed;filaments ca. 1 mm. Female flowers: corolla larger than male
flowers, 12–15 mm in diam.; ovary ovoid, ca. 10 × 5 mm. Fruitellipsoid, 3.5–4 × ca. 2.5 cm, base obtuse-rounded; fruit wall

Tuber oblate-ovoid. Stem and branchlets slightly stout,
distinctly sulcate. Tendrils 12–25 cm. Leaves pedately 5–7foliolate; petiole 3–6 cm; leaflets lanceolate to obovate-lanceolate, margin crenate; median leaflets ca. 14 × 1.5–4 cm; petiolule 5–10 mm. Male inflorescence dichotomously cymose; peduncle flexuous, 5–12 cm, densely pubescent; calyx segments
ovate-triangular, ca. 5 × 3 mm, apex reflexed; corolla paleyellow-green, bowl-like, ca. 10 mm in diam.; segments oblong,
7–8 × 3–4 mm; filaments ca. 1 mm. Female flowers unknown.
Fruit clavate or clavate-elliptic, 7–8 × 2–2.5 cm, distinctly
costate; fruiting pedicel 10–15 mm. Seeds dark brown, elliptic,
ca. 12 × 7 mm, almost unwinged, margin 1–3 mm wide. Fl.
Jul–Oct, fr. Sep–Nov.
● Broad-leaved forests, forest margins. SW Yunnan.

19. Hemsleya zhejiangensis C. Z. Zheng, Acta Phytotax. Sin.
23: 67. 1985.
浙江雪胆 zhe jiang xue dan
Tuber enlarged. Stem and branchlets slender, sulcate,nodes densely pubescent. Leaves pedately 5–9-foliolate; petiole
2.5–5 cm; leaflets elliptic-lanceolate, margin sparsely dentate;
median leaflets 6–11 × 2–3.5 cm. Male inflorescence dichotomously cymose; peduncle flexuous, 13–17 cm; calyx segments
ovate-lanceolate, 4–5 × 3–4 mm, apex reflexed; corolla pale

CUCURBITACEAE

9yellow-green, bowl-like, 8–10 mm; segments obovate, 7–8 ×
4–5 mm, fleshy, apex reflexed; filaments ca. 1 mm. Female
flower unknown. Fruit broadly to narrowly elliptic, (6–)11–17
× 2–3 cm, indistinctly costate; fruiting pedicel with distinct
articulation, 1–2 cm. Seeds dark brown, oblong, almost
unwinged, margin 2–4 mm. Fl. Jun–Sep, fr. Aug–Nov.

20b. Hemsleya chinensis var. ningnanensis L. D. Shen & W.
J. Chang, Acta Phytotax. Sin. 21: 185. 1983.

● Broad-leaved forests, bamboo forests; 800–1000 m. Anhui,
Jiangxi, Zhejiang.

Branchlets, petiole, and fruit without spiculiform hairs.
Corolla 10–12 mm in diam., usually with dense white hairs.
Fruit obovoid; pedicel with articulation.

20. Hemsleya chinensis Cogniaux ex F. B. Forbes & Hemsley,
J. Linn. Soc., Bot. 23: 490. 1888.
雪胆 xue dan
Tuber ovoid or oblate. Stem and branchlets slender. Tendrils 8–14 cm. Leaves pedately 5–9-foliolate; petiole 4–8 cm;
leaflets ovate-lanceolate, oblong-lanceolate, or broadly lanceolate, margin crenate; median leaflets 5–12 × 2–2.5 cm; petiolule
5–10 mm. Male inflorescence dichotomously cymose or paniculate; peduncle filiform, flexuous, 5–12 cm; calyx segments
ovate, ca. 7 × 4–5 mm, apex acute, reflexed; corolla orange-red,
Chinese lantern-shaped or rounded-globose, 8–15 mm in diam.;segments oblong, 10–13 × 8–9 mm, subglabrous; filaments ca.
1 mm. Female inflorescence simple; peduncle 2–4 cm; calyx
and corolla as in male flowers; corolla ca. 15 mm in diam.;ovary cylindric, 5–10 × 2–3 mm. Fruit ellipsoid, 3–7 × ca. 2
cm, base attenuate; fruiting pedicel slightly curved, with orwithout articulation, 8–10 mm. Seeds brown, suborbicular, 10–
12 × ca. 10 mm, ca. 3 mm thick, unwinged, margin ca. 1 mm
wide. Fl. Jul–Sep, fr. Sep–Nov. 2n = 28*.
● Evergreen broad-leaved forests, forest margins; 400–2800 m.
Guizhou, Hubei, Sichuan, Yunnan.

1a. Fruit with white spiculiform hairs along
longitudinal ribs; branchlets and petiole
with dense spiculiform hairs ............... 20d. var. polytricha
1b. Fruit without spiculiform hairs along
longitudinal ribs; branchlets and petiole
without spiculiform hairs.
2a. Corolla usually glabrous,
12–15 mm in diam.; fruit
elliptic, 3–7 cm ............................. 20a. var. chinensis
2b. Corolla densely white pubescent,
8–12 mm in diam.
3a. Corolla 10–12 mm in diam.;
fruit obovoid, pedicel
with articulation .............. 20b. var. ningnanensis
3b. Corolla ca. 8 mm in diam.;
fruit ovoid, pedicel
without articulation .......... 20c. var. longevillosa
20a. Hemsleya chinensis var. chinensis
雪胆(原变种) xue dan (yuan bian zhong)Branchlets, petiole, and fruit without spiculiform hairs.
Corolla 12–15 mm in diam., usually glabrous. Fruit elliptic, 3–7cm.
● Evergreen broad-leaved forests, forest margins; 400–2800 m.
Guizhou, Hubei, Sichuan, Yunnan.

宁南雪胆 ning nan xue dan
Hemsleya ningnanensis L. D. Shen & W. J. Chang; H.
villosipetala C. Y. Wu & Z. L. Chen.

● Evergreen broad-leaved forests. SW Sichuan, NE Yunnan.
Hemsleya chinensis var. ningnanensis is not a new combination
based on H. ningnanensis (L. D. Shen & W. J. Chang, Acta Acad. Med.
Sichuan. 11(1): 19. 1980), although both names have the same type.

20c. Hemsleya chinensis var. longevillosa (C. Y. Wu & Z. L.
Chen) D. Z. Li, Syst. Evol. Hemsleya, 95. 1993 [“longivillosa”].
长毛雪胆 chang mao xue dan
Hemsleya longevillosa C. Y. Wu & Z. L. Chen, Acta
Phytotax. Sin. 23: 140. 1985.
Branchlets, petiole, and fruit without spiculiform hairs.
Corolla usually with dense white hairs, ca. 8 mm in diam. Fruitovoid; pedicel without articulation.
● Evergreen broad-leaved forests; 1900–2100 m. Yunnan (Fuyuan).

20d. Hemsleya chinensis var. polytricha Kuang & A. M. Lu,
Acta Phytotax. Sin. 20: 90. 1982.
毛雪胆 mao xue dan
Branchlets and petiole with dense spiculiform hairs. Fruitwith dense spiculiform hairs along longitudinal ribs.
● Broad-leaved forests; 1300–1500 m. W Hubei.

21. Hemsleya gigantha W. J. Chang, Acta Phytotax. Sin. 21:
186. 1983.
巨花雪胆 ju hua xue dan
Tuber yellow-brown. Leaves pedately 7–9-foliolate; petiole 5–9 cm; leaflets broadly elliptic-lanceolate or ovate-lanceolate, base cuneate, margin crenate; median leaflets 7–12 × 3–5
cm. Male inflorescence dichotomously cymose; rachis 7–11
cm; calyx segments lanceolate, ca. 15 × 7 mm, apex reflexed;
corolla orange-red, Chinese lanternlike, 1.5–2(–2.5) cm indiam.; segments broadly ovate, 20–25 × 15–18 mm, revolute,obtuse-rounded and apiculate at apex; filaments ca. 1 mm. Female inflorescence simple; ovary ellipsoid, 10–15 × 6–10 mm.
Fruit subspheroid to ovoid, 2–3.5 cm in diam., fruit wall leathery. Seeds dark brown, broadly ovoid, 10–12 × 8–10 mm, 2–3
mm thick, almost unwinged; margin narrow, ca. 2 mm wide,slightly erose. Fl. Jun–Sep, fr. Aug–Nov.
● Broad-leaved or Pinus forests, forest margins. SW Sichuan.

22. Hemsleya dolichocarpa W. J. Chang, Acta Phytotax. Sin.
21: 190. 1983.
长果雪胆 chang guo xue dan

CUCURBITACEAE

10Tuber enlarged, yellow-brown. Stem and branchlets slender. Leaves pedately 5–7-foliolate; petiole 3.5–6.5 cm; leaflets
obovate-lanceolate or elliptic-lanceolate, margin crenate; median leaflets 7–15 × 4–7 cm. Male inflorescence dichotomously
cymose; rachis 3–8 cm; calyx segments lanceolate, 6–8 × 4–6
mm, apex apiculate, flexuous; filaments ca. 1 mm. Female inflorescence simple; ovary cylindric, ca. 6 × 3 mm. Fruit elliptic,
5–8 × 2–3.5 cm. Seeds broadly ovate to suborbicular, compressed, 13–14 × ca. 11 mm, 2–3 mm thick, unwinged; marginca. 1 mm wide, erose. Fl. Jun–Sep, fr. Aug–Nov.
● Broad-leaved forests; 1400–1800 m. C and SW Sichuan.

23. Hemsleya macrosperma C. Y. Wu, Acta Phytotax. Sin. 23:
139. 1985.
罗锅底 luo guo di
Tuber enlarged, oblate. Stem and branchlets slender. Tendrils 6–10 cm. Leaves pedately 5–7-foliolate; petiole 4–5 cm;
leaflets oblong-lanceolate to obovate-lanceolate, margin crenate;
median leaflets 4–9 × 1.5–3 cm; petiolule 2–4 mm. Male inflorescence dichotomously cymose; peduncle 2–8 cm; calyx segments ovate, 4–5 × ca. 3 mm, apex acute, reflexed; corollaorange-red, bowl-like, 8–10 mm in diam.; segments oblong, ca.
8 × 4 mm, sparsely white villous; filaments ca. 2 mm. Female
inflorescence: peduncle 1–5 cm; ovary ellipsoid or subglobose,
5–7 × 4–5 mm. Fruit ovoid, 3.5–5 cm in diam. Seeds darkbrown, ovoid, 9–11 × 8–9 mm, unwinged, margin smooth, ca. 1
mm wide. Fl. Jul–Sep, fr. Sep–Nov.
● Evergreen broad-leaved or Pinus forests; 1800–3200 m. SW Sichuan, C and NE Yunnan.

1a. Fruit ovoid, 3.5–4 cm ..................... 23a. var. macrosperma
1b. Fruit elliptic, to 5 cm ...................... 23b. var. oblongicarpa
23a. Hemsleya macrosperma var. macrosperma
罗锅底(原变种) luo guo di (yuan bian zhong)Fruit ovoid, 3.5–4 cm.

● Evergreen broad-leaved or Pinus forests; 1800–3200 m. SW Sichuan, C and NE Yunnan.

23b. Hemsleya macrosperma var. oblongicarpa C. Y. Wu &Z. L. Chen, Acta Phytotax. Sin. 23: 140. 1985.
长果罗锅底 chang guo luo guo di
Fruit elliptic, to 5 cm.
● Evergreen broad-leaved forests; 1900–2000 m. SW Sichuan,
NE Yunnan.

24. Hemsleya dulongjiangensis C. Y. Wu, Acta Phytotax. Sin.
23: 134. 1985.
独龙江雪胆 du long jiang xue dan
Tuber enlarged. Stem and branchlets slender, longitudinally sulcate, densely pubescent. Tendrils 6–9 cm, densely pubescent. Leaves pedately 3–7-foliolate; petiole 2–3 cm; leaflets
broadly ovate-lanceolate to rhombic, margin sparsely serrate;
median leaflets 3–6 × 3–3.5 cm. Flowers unknown. Fruit ovoid,
3.5–4.5 × 3–3.5 cm; fruiting pedicel curved, 5–8 cm. Seeds(immature) yellow-brown, broadly ovoid, ca. 13 mm in diam.,
unwinged; margin 1–2 mm wide, erose. Fr. Sep.
● Open forests in valleys; 1400–2700 m. NW Yunnan.

25. Hemsleya endecaphylla C. Y. Wu, Acta Phytotax. Sin. 23:
142. 1985.
十一叶雪胆 shi yi ye xue dan
Tuber enlarged. Stem slightly robust. Tendrils 8–12 cm.
Leaves pedately 9–11-foliolate; petiole stout, 5–7 cm; leaflets
obovate-lanceolate, margin irregularly crenate; median leaflets
5–7 × 2–2.5 cm; petiolule 3–5 mm. Male inflorescence paniculate; rachis indistinctly flexuous, 5–8 cm; calyx segments paleyellow-green, ovate-triangular, ca. 5 × 4 mm, apex reflexed; corolla orange-yellow, bowl-like, 8–10 mm in diam.; segments
oblong, with dense squamose hairs, apex reflexed; filaments
very short, ca. 0.5 mm. Female flower and fruit unknown. Fl.
Jun–Jul.
● Broad-leaved forests; ca. 2400 m. NW Yunnan.

2. GOMPHOGYNE Griffith, Account Bot. Coll. Cantor, 26. 1845.
锥形果属 zhui xing guo shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, climbing; stems slender, angular. Leaves petiolate, pedately 7–9-foliolate; leaflet blade lanceolate or oblong, dentate.
Tendrils 2-fid. Plants monoecious. Flowers small, pale green. Male flowers in racemes or panicles; calyx rotate, 5-partite; segments
oblong-lanceolate, margins erose; corolla rotate, 5-partite; segments oblong-lanceolate, erose, caudate-acuminate; stamens 5; filaments short, united at base; anthers erect, subglobose, 1-celled, dehiscence longitudinal; rudimentary ovary absent. Female flowers inpanicles or fascicled in axils of leaves; calyx and corolla as in male flowers; ovary clavate, 1-locular; styles 3; stigmas 2-lobed;
ovules 3, pendulous from apex of locules; staminodes absent. Fruit capsular, turbinate, venose and ribbed, foveolate, truncate at apex,
3-valved, crowned by persistent styles. Seeds 3, black, oblong, testa thick, margin thickened, erose.
Two species: Bhutan, China, India, Nepal; one species in China.

1. Gomphogyne cissiformis Griffith, Account Bot. Coll. Cantor, 26. 1845.
锥形果 zhui xing guo

Stem glabrous or slightly puberulent. Leaves membranous;
petiole 3–6 cm, glabrous; leaflets obovate-oblong, glabrous,
median leaflets 4–6 × 1.5–2 cm, lateral leaflets shorter, base attenuate, margin crenate-dentate, apex acuminate or acute; petio-

CUCURBITACEAE

lules 3–9 cm. Tendrils slender, 2-fid. Plants monoecious. Male
flowers in racemes or panicles; rachis slender, flexuous, 10–30
cm, glabrous; pedicels capillary-like, often fasciculate, 1–3 mm,with bracteoles at base; calyx segments narrowly ovate, 1–1.5
mm, acute; corolla segments ovate-lanceolate, 2.5–3 × ca. 1
mm, 3-veined, entire or indistinctly dentate. Female flowers inpanicles 5–12 cm, or fascicled in axils of leaves; pedicels filiform, reflexed; bracteoles subulate; ovary funnelform. Fruit palegreen, turbinate, 12–16 × 8–11 mm, apex truncate, 3-longitudinally dehiscent, 1–3-seeded. Seeds dark brown, oblong, 7(–12)
× 3–4 mm, 1–2 mm thick, verrucose, base acute, margin dentate, apex truncate.
Forests on mountainsides; 2100–2800 m. S and SW Yunnan [Bhutan, India, Nepal].

1a. Fruit glabrous ........................................ 1a. var. cissiformis
1b. Fruit tomentose ........................................... 1b. var. villosa

11

1a. Gomphogyne cissiformis var. cissiformis
锥形果(原变种) zhui xing guo (yuan bian zhong)Fruit glabrous.
Forests on mountainsides; 2100–2800 m. S and SW Yunnan [Bhutan, India, Nepal].

1b. Gomphogyne cissiformis var. villosa Cogniaux in A. Candolle & C. Candolle, Monogr. Phan. 3: 925. 1881.
毛锥形果 mao zhui xing guo
Gomphogyne cissiformis f. villosa (Cogniaux) M. Mizushima.
Fruit tomentose.
Forests on mountainsides; ca. 2300 m. S and SW Yunnan (Menghai, Yongde) [NE India (Sikkim), Nepal].

3. GYNOSTEMMA Blume, Bijdr. 23. 1825.
绞股蓝属 jiao gu lan shu
Chen Shukun (陈书坤), Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Enkylia Griffith; Pestalozzia Zollinger & Moritzi; Trirostellum Z. P. Wang & Q. Z. Xie.
Herbs, perennial, scandent, glabrous or pubescent. Leaves alternate, pedate, with 3–9 leaflets, rarely simple; leaflet blade ovatelanceolate. Tendrils bifurcate, rarely simple. Plants dioecious, rarely monoecious. Flowers unisexual, in racemes or panicles, axillary
or terminal; pedicel articulate; bracteole basal. Male flower: calyx tube short, 5-lobed; segments narrowly lanceolate; corolla
greenish or white, rotate, deeply 5-lobed; segments lanceolate or ovate-lanceolate, involute in bud; stamens 5, inserted at base of
perianth tube; filaments short, connate; anthers erect, ovoid, 2-celled, dehiscence longitudinal; connective narrow, but not extended;pollen grains globose or elliptic, longitudinally striate or smooth, dehiscent by pores; rudimentary pistil absent. Female flowers:
calyx and corolla as in male flower; staminodes present; ovary globose, 2–5-locular; styles 3, rarely 2, 4, or 5, free; stigmas 2 or 1
and crescent-shaped and irregularly incised-dentate; ovules 2, pendulous in each locule. Berry globose, in form and size like a pea, or
capsule, 3-lobed from top, top umbonulate or with 3 persistent long styles. Seeds 2 or 3, broadly ovate, compressed, with papillae orspiniform papillae.
About 17 species: tropical Asia to E Asia, from the Himalaya to Japan, Malaysia, and New Guinea; 14 species (nine endemic) in China.

1a. Capsule campanulate, dehiscent when mature along ventral suture, 3-lobed; persistent styles 3(–5), long
rostriform.
2a. Female flowers solitary or paired, axillary, pedicel 3–4 cm; ovary 5-locular, styles 5 .................................... 1. G. pentagynum
2b. Female flowers numerous, in racemes or panicles, axillary or terminal, pedicel shorter than 5 mm;ovary 3-locular, styles 3.
3a. Female flowers in lax panicle; fruit stalk 8–10 mm ...................................................................................... 6. G. laxiflorum
3b. Female flowers in spiciform racemes; fruit stalk shorter than 5 mm.
4a. Stigma crescent-shaped and irregularly incised-dentate, style 2.5–3 mm, or thick and short,
ca. 0.5 mm.
5a. Styles thin and long, 2.5–3 mm; persistent styles up to 5 mm; side of seed not sulcate andwithout wing ..................................................................................................................................... 2. G. yixingense
5b. Styles short and thick; persistent styles shortly rostriform; seed broadly cordate, margin
sulcate and with narrow wing ................................................................................................... 3. G. cardiospermum
4b. Stigma 2-lobed, forked, never crescent-shaped, style shorter than 0.5 mm.
6a. Fruit ca. 3 mm in diam., glabrous, dark punctate; leaves with 5 elliptic leaflets ..................... 4. G. microspermum
6b. Fruit 5–6 mm in diam., white villous, without dark spots; leaves with 5–7
obovate-elliptic leaflets .................................................................................................................. 5. G. aggregatum
1b. Berry globose or depressed-globose, not dehiscent; persistent styles short, umbonate, never rostrate.
7a. Leaves simple ................................................................................................................................................. 7. G. simplicifolium
7b. Leaves compound, pedate.
8a. Stem 3-angled, with narrow wings .......................................................................................................... 14. G. caulopterum
8b. Stem cylindrical, with longitudinal striae, but without wings.

CUCURBITACEAE

129a. Leaves with 3–5 leaflets.
10a. Leaf smooth, glabrous on both surfaces, or with hairs only along midrib; stem with only
sparse hairs at nodes; corolla segments narrowly lanceolate, 2–3 mm ................................................ 8. G. laxum
10b. Leaf blade and stem all densely pubescent; corolla segments oblong-elliptic, ca. 2 mm .... 9. G. “burmanicum”
9b. Leaves with (3–)5–7(–9) leaflets.
11a. Fruit densely hirsute-pubescent; stem stout ............................................................................ 10. G. pentaphyllum
11b. Fruit glabrous; stem slender.
12a. Fruit stalk shorter than 5 mm; leaves sparsely to densely pubescent ........................... 10. G. pentaphyllum
12b. Fruit stalk 7–25 mm.
13a. Plants monoecious; fruit 3-angled-obovoid or 3-angled-oblate ............................ 11. G. guangxiense
13b. Plants dioecious; fruit globose or obtriangular.
14a. Fruit obtriangular, compressed; leaflets 5–7, elliptic or elliptic-ovate ......... 12. G. compressum
14b. Fruit globose; leaflets 7–9, rhombic-elliptic or obovate-lanceolate ................... 13. G. longipes
1. Gynostemma pentagynum Z. P. Wang, Acta Bot. Yunnan.
11: 165. 1989.
五柱绞股蓝 wu zhu jiao gu lan
Plants with short and succulent rootstocks. Stem ca. 2 m or
more, ca. 4 mm in diam., angular, white villous. Tendrils filiform, villous below, gradually glabrescent above, apex bifid.
Leaves pedate, with (5–)7 leaflets; petiole 9–13 cm, villous;
leaflet blade elliptic, central leaflet ca. 10 cm, both surfaces
sparsely pubescent, densely villous along veins, margin irregularly serrate, apex shortly acuminate; petiolule 3–5 mm; lateral
leaflet blades gradually smaller and asymmetrical, petiolule
shorter; outermost leaflet without petiolule. Plants dioecious.
Male flowers numerous in panicle, 3–4 cm, villous; calyx segments narrowly elliptic, ca. 0.9 × 0.5 mm, both surfaces glabrous, apex obtuse; corolla segments ovate, ca. 4 × 0.6 mm,outside glabrous, inside glandular hairy, with obscure midrib,
apex filiform-acuminate; filaments ca. 0.2 mm, connective
cylindrical. Female flower(s) solitary or 2(or 3) in very shortraceme; peduncles 2–3 mm; pedicels up to 4 cm; calyx and corolla as in male flowers; staminodes absent; gynoecium with (4or)5 carpels; ovary densely pubescent and glandular pubescent,(4 or)5-locular, each locule with 1 ovule; styles (4 or)5, divaricate; stigmas 2, explanate. Fruit immature. Fl. Jul.
● Evergreen broad-leaved forests, moist mountains. NW Hunan(Dayong).

2. Gynostemma yixingense (Z. P. Wang & Q. Z. Xie) C. Y. Wu& S. K. Chen, Acta Phytotax. Sin. 21: 364. 1983.
喙果绞股蓝 hui guo jiao gu lan
Trirostellum yixingense Z. P. Wang & Q. Z. Xie, Acta
Phytotax. Sin. 19: 483. 1981; T. yixingense var. trichocarpum J.
N. Ding.
Plants to over 10 m. Stem slender, angular-sulcate, villous
only at nodes. Leaves membranous, pedately 5–7-foliolate;petiole 3–6 cm; leaflets elliptic, median leaflets 4–8 cm, lateral
leaflets smaller, veins pubescent, base cuneate, margin crenate,apex acuminate; petiolule ca. 5 mm. Tendrils filiform, simple.
Male flowers in panicle; rachis 9–12 cm, sparsely pubescent;calyx segments elliptic-lanceolate, 1–1.5 × ca. 0.5 mm, apex
obtuse; corolla pale green; segments ovate-lanceolate, 2–2.5mm, caudate-acuminate; filaments connected; anthers ca. 0.2
mm. Female flowers fasciculate in axils of leaves; calyx andcorolla as in male flowers; ovary subglobose, 1.5–2 mm indiam., sparsely puberulent; styles 3, 2.5–3 mm; stigmas halfmoon-shaped, margin dentate; staminodes 5, subulate. Capsulecampanulate, ca. 5 × 8 mm, glabrous, apex truncate. Seeds
broadly cordate, ca. 3 × 4 mm. Fl. Aug–Sep, fr. Sep–Oct.
● Forests, thickets; below 100 m. Anhui, S Jiangsu, Zhejiang.
One of us (Jeffrey) treats Gynostemma laxiflorum as a synonym of
this species.

3. Gynostemma cardiospermum Cogniaux ex Oliver,
Hooker’s Icon. Pl. 23: t. 2225. 1892 [“cardiosperma”].
心籽绞股蓝 xin zi jiao gu lan
Trirostellum cardiospermum (Cogniaux ex Oliver) Z. P.
Wang & Q. Z. Xie.
Herbs, scandent. Stem slender, angular-sulcate, glabrous.
Leaves pedately 3–7-foliolate; petiole 2.5–5 cm; leaflets lanceolate or oblong-elliptic, median leaflets 4–10 cm, lateral leaflets
shorter, glabrous or setose only on veins, base attenuate, margin
irregularly crenate, apex acuminate. Tendrils slender, 2-fid. Male
flowers in a panicle; rachis slender; calyx segments oblonglanceolate, acute; corolla segments lanceolate, 1-veined, caudate-acuminate; anthers ovoid, 1-celled. Female flowers in a
raceme; calyx and corolla as in male flowers; ovary globose,
villous; ovules 2; styles 3, ca. 0.5 mm; stigmas half-moonshaped, margin dentate. Capsule globose or subcampanulate,ca. 8 mm in diam., apex truncate. Seeds broadly cordate, 4.2–5
mm in diam., slightly compressed, both surfaces rugose-verrucose, margin narrowly winged. Fl. Jun–Aug, fr. Aug–Oct.
● Forests and thickets on mountain slopes; (1400–)1900–2300 m.
W Hubei, S Shaanxi, Sichuan.

4. Gynostemma microspermum C. Y. Wu & S. K. Chen, Acta
Phytotax. Sin. 21: 364. 1983 [“microsperma”].
小籽绞股蓝 xiao zi jiao gu lan
Stem slender, angular-sulcate, glabrous or pubescent at
nodes. Leaves pedately 5-foliolate; petiole ca. 2 cm, glabrous;
leaflets elliptic, median leaflets 3–4 × 1.5–2 cm, lateral leaflets
smaller, thinly papery, adaxially pubescent, base obtuse, margin
irregularly crenate, apex obtuse, cuspidate. Tendrils filiform, 2fid, glabrous. Flowers unknown. Infructescence racemose,dense. Capsule yellow-green, dark punctate, globose, ca. 3 mm
in diam., glabrous, apex truncate, dehiscent, 1- or 2-seeded.

CUCURBITACEAE

Seeds brown, broadly cordate, compressed, ca. 2.5 × 2 mm, lessthan 1 mm thick, papillose. Fr. Dec.
Dense forests; 800–1400 m. S Yunnan [N Thailand].

5. Gynostemma aggregatum C. Y. Wu & S. K. Chen, Acta
Phytotax. Sin. 21: 365. 1983.
聚果绞股蓝 ju guo jiao gu lan
Stem slender, angular-sulcate, glabrous or pubescent at
nodes. Leaves pedately 5–7-foliolate; petiole 2–2.5 cm, glabrous; leaflets obovate-elliptic, median leaflets 3–3.5 × ca. 1.5
cm, lateral leaflets smaller, membranous, abaxially glabrous,
adaxially sparsely pubescent, lateral veins 6–9 pairs, base obtuse, margin crenate, apex obtuse-rounded, cuspidate; petiolule
2–5 mm, glabrous. Tendrils filiform, 2-fid. Male flowers unknown. Female flowers in a small 3- or 4-flowered panicle;
bracts capillary; calyx segments very small, subulate; corollasegments lanceolate, ca. 1 mm, apex shortly acuminate; ovary
globose, ca. 1 mm in diam., villous; styles 3, free, ca. 0.75 mm;stigma 2-lobed. Capsules aggregated in a dense 1.5–2 cm spike;fruit broadly campanulate, ca. 5 mm in diam., white pubescent,apex truncate. Seeds gray-brown, ovoid, ca. 3 × 2 mm, ca. 1
mm thick, verruculose, margin dentate. Fl. Jul–Aug, fr. Sep–
Oct.
● Pinus forests, mixed forests; 2300–2700 m. NW Yunnan.

6. Gynostemma laxiflorum C. Y. Wu & S. K. Chen, Acta
Phytotax. Sin. 21: 366. 1983.
疏花绞股蓝 shu hua jiao gu lan
Stem slender, angular-sulcate, glabrous or puberulent at
nodes. Leaves pedately 5-foliolate; petiole ca. 3 cm; leaflets
oblong-lanceolate, median leaflets ca. 4 × 1.5 cm, lateral leaflets smaller, membranous, abaxially glabrous, adaxially sparsely
puberulent, base obtuse, margin dentate, apex acuminate. Tendrils simple, glabrous. Male flowers unknown. Female flowers:
inflorescence a sparse panicle, axillary or terminal, 2–5 × 1.5–
2.5 cm, rachis and branches glabrous; pedicel filiform, 4–5 mm;bracts linear; calyx segments oblong-lanceolate, ca. 1.75 × 0.5mm, glabrous, apex shortly acuminate; corolla segments ovatelanceolate, ca. 2.5 × 1 mm, puberulent or glabrescent, apex caudate-acuminate; staminodes 5, terete; ovary globose, ca. 2 mm
in diam., pubescent; styles 3, ca. 1.5 mm, base puberulent;
stigma luniform, irregularly 3–5-dentate. Fruiting pedicel 8–10mm, glabrous; capsule campanulate, 6–7 mm, glabrous, apex
truncate, 3-rostrate, beak 2–3 mm, dehiscent, 2- or 3-seeded.
Seeds brown, broadly cordate, ca. 2 × 2.5 mm, ca. 1.5 mm
thick, papillate. Fl. and fr. Nov–Dec.
● Mountain slopes; below 300 m. S Anhui.
One of us (Jeffrey) treats this taxon as a synonym of Gynostemma
yixingense.

7. Gynostemma simplicifolium Blume, Bijdr. 24. 1825 [“simplicifolia”].
单叶绞股蓝 dan ye jiao gu lan
Gynostemma pentaphyllum (Thunberg) Makino f. simplicifolium (Blume) W. J. de Wilde & Duyfjes.

13

Stem slender, branched, angular-sulcate, pubescent. Leaves
simple; leaf blade ovate, 10–15 × 8–9 cm, papery, abaxially glabrous, adaxially puberulent only on veins, basal veins 7–9, baserounded to slightly cordate, margin crenate, apex acuminate;petiole 4–6 cm, pubescent. Tendrils 2-fid. Male flowers: inflorescence paniculate; peduncle filiform, 10–25 cm, pubescent;pedicels filiform, 0.5–2 mm; calyx segments oblong-lanceolate,
0.5–1 mm, puberulent, apex obtuse; corolla pale green-white orpale green-yellow; segments oblong, ca. 3 × 0.7–1 mm, 1veined, apex obtuse. Fruit pale yellow-green, black when mature, globose, 7–8 mm in diam. Seeds broadly ovate, ca. 4 × 3.5mm, compressed, verrucose, base rounded, apex acute.
Forests; 1300–1400 m. Hainan, S Yunnan [India, Indonesia, Malaysia, Myanmar, Philippines].
One of us (Jeffrey) treats this taxon as a synonym of Gynostemma
pentaphyllum.

8. Gynostemma laxum (Wallich) Cogniaux in A. Candolle &C. Candolle, Monogr. Phan. 3: 914. 1881.
光叶绞股蓝 guang ye jiao gu lan
Zanonia laxa Wallich, Pl. Asiat. Rar. 2: 29. 1831; Alsomitra laxa (Wallich) M. Roemer; Gynostemma crenulatum Ridley; Pestalozzia laxa (Wallich) Thwaites; Z. wightiana Arnott.
Stem slender, much branched, glabrous or sparsely puberulent. Leaves pedately 3-foliolate; petiole 1.5–4 cm, glabrous; median leaflet oblong-lanceolate, sometimes rhombic,
5–10 × 2–3 cm, papery, base broadly cuneate, apex acute or
shortly acuminate; lateral leaflets ovate, 4–7 × 2–3.5 cm,
glabrous, margin undulate-crenate; petiolule (2–)5–7 mm. Male
flowers: inflorescence paniculate, terminal or axillary, (5–)10–
30 cm, pubescent; bracts subulate-lanceolate, ca. 1 mm; pedicels filiform, 3–7 mm; bracteole subulate, small; calyx segments narrowly triangular-ovate, ca. 0.5 mm; corolla yellowgreen; segments narrowly ovate-lanceolate, ca. 1.5 × 0.5 mm,
glabrous, 1-veined, entire, acuminate at apex. Female inflorescence as male flowers; corolla segments narrowly triangular;
ovary globose, ca. 1 mm in diam.; styles 3, free, apex 2-fid.
Fruit yellow-green, globose, 8–10 mm, glabrous, indehiscent.
Seeds pale gray, broadly ovate, ca. 4 mm in diam., compressed,
both surfaces papillate, base rounded, apex acute. Fl. Aug, fr.
Aug–Sep.
Dense forests in valleys. Guangxi, Hainan, SE Yunnan [India, Indonesia, Malaysia, Myanmar, Nepal, Philippines, Thailand, Vietnam].
One of us (Jeffrey) would treat this taxon as a synonym of Gynostemma pentaphyllum.

9. Gynostemma “burmanicum” King ex Chakravarty, Indian
J. Agric. Sci. 16(1): 86. 1946 [“Gymnostemma burmanica”].
缅甸绞股蓝 mian dian jiao gu lan
Stem angular-sulcate, densely pubescent at nodes. Leaves
papery, 3-foliolate, both surfaces hispidulous; petiole 3–6.5 cm,
densely pubescent; median leaflet rhombic, (6–)8–12 × 3–5.5
cm, base broadly cuneate, margin crenate, apex shortly acuminate; lateral leaflets 4–9 × ca. 4 cm, outside subovate, baserounded, inside sublanceolate or oblanceolate, base attenuate,
lateral veins 8 or 9; median petiolule 5–6 mm; lateral leaflets

CUCURBITACEAE

14subsessile. Tendrils 2-fid. Male flowers in panicle; calyx segments oblong, ca. 0.75 × 0.3 mm, apex obtuse; corolla green;
segments ± elliptic, apex acute or shortly acuminate; stamens
ca. 1 mm, glabrous. Female flowers unknown. Fruit green, baccate, globose, 5–10 mm in diam., glabrous, 3-seeded. Seeds
pale brown, broadly ovate, ca. 3(–5) × 3 mm, ca. 2 mm thick,compressed, both surfaces papillate, base subrounded, edges
with furrow.
Open forests or thickets, forests in mountain valleys; 600–1300 m.
S and SE Yunnan [Myanmar, Thailand].
One of us (Jeffrey) considers this species and its varieties, thenames of which have never been validly published, to represent densely
hairy variants of Gynostemma pentaphyllum. The name G. “burmanicum” was not validly published by Chakravarty (loc. cit.) because no
Latin description or diagnosis was provided (Vienna Code, Art. 36.1); asa result, the name G. “burmanicum var. molle” was not validly published either (Art. 43.1). A subsequent publication of the species name
(Chakravarty, Rec. Bot. Surv. India 17(1): 190. 1959) again failed toprovide a Latin description or diagnosis and did not indicate a type.

1a. Fruit 5–6 mm in diam.; seeds
3(–3.5) mm .................................... 9a. var. “burmanicum”
1b. Fruit 8–10 mm in diam.; seeds
ca. 5 mm .................................................... 9b. var. “molle”
9a. Gynostemma “burmanicum var. burmanicum”
缅甸绞股蓝(原变种) mian dian jiao gu lan (yuan bian zhong)Fruit 5–6 mm in diam.; seeds 3(–3.5) mm.
Open forests and thickets; 800–1200 m. S Yunnan [Myanmar,
Thailand].

9b. Gynostemma “burmanicum var. molle” (C. Y. Wu, Acta
Phytotax. Sin. 21: 360. 1983).
大果绞股蓝 da guo jiao gu lan
Fruit 8–10 mm in diam.; seeds ca. 5 mm.
● Forests in mountain valleys or open forests; 600–1300 m. S andSE Yunnan.

10. Gynostemma pentaphyllum (Thunberg) Makino, Bot.
Mag. (Tokyo) 16: 179. 1902.
绞股蓝 jiao gu lan
Stem and branches slender, angular-sulcate, glabrous orsparsely pubescent. Leaves pedately 3–9-foliolate, usually 5–7foliolate, membranous or papery, pubescent or glabrous; leaflets
ovate-oblong or lanceolate, median leaflets 3–12 × 1.5–4 cm,
lateral leaflets smaller, both surfaces sometimes hispid, lateralveins 6–8 pairs, base attenuate, margin crenate, apex acute or
shortly acuminate; petiolule 1–5 mm. Tendrils filiform, 2-fid.
Male flowers in panicle; peduncle filiform, 10–15(–30) cm,
many branched; pedicels filiform, 1–4 mm; bracteole subulate;calyx tube very short; segments triangular, ca. 0.7 mm, apex
acute; corolla pale green or white; segments ovate-lanceolate,
2.5–3 × ca. 1 cm, 1-veined, apex long acuminate. Female flowers: panicle shorter than that of male flowers; calyx and corolla
as in male flowers; ovary globose, 2- or 3-loculed; styles 3;
stigmas 2-lobed; staminodes 5, short. Fruit indehiscent, blackwhen mature, globose, 5–6 mm in diam., 2-seeded, glabrous ordensely hispid and pubescent. Seeds brown, ovate-cordate, ca. 4

mm in diam., compressed, both surfaces papillose, base cordate,
apex obtuse. Fl. Mar–Nov, fr. Apr–Dec.
Forests, thickets or roadsides on mountain slopes; 300–3200 m.
Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, S Henan,
Hubei, Hunan, Jiangsu, Jiangxi, Shandong, Sichuan, Taiwan, Yunnan,
Zhejiang [Bangladesh, Bhutan, India, Indonesia, S Japan, S Korea,
Laos, Malaysia, Myanmar, Nepal, New Guinea, Sri Lanka, Thailand,
Vietnam].

1a. Fruit glabrous .................................. 10a. var. pentaphyllum
1b. Fruit densely hispid andpubescent .......................................... 10b. var. dasycarpum
10a. Gynostemma pentaphyllum var. pentaphyllum
绞股蓝(原变种) jiao gu lan (yuan bian zhong)Vitis pentaphylla Thunberg in Murray, Syst. Veg., ed. 14,244. 1784; Alsomitra cissoides M. Roemer; Enkylia digyna
Griffith; E. trigyna Griffith; Gomphogyne alleizettei Gagnepain;
Gynostemma pallidinerve Z. Zhang; G. pedatum Blume; G.
pedatum var. hupehense Pampanini; G. pedatum var. pubescens
Gagnepain; G. pedatum var. trifoliatum Hayata; G. pubescens(Gagnepain) C. Y. Wu; G. siamicum Craib; G. zhejiangense X. J.
Xue; Pestalozzia pedata (Blume) Zollinger & Moritzi; V. mairei
H. Léveillé (1912), not H. Léveillé (1909); V. martini H.
Léveillé & Vaniot; Zanonia pedata (Blume) Miquel.
Fruit glabrous.
Forests, thickets or roadsides on mountain slopes; 300–3200 m.
Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, S Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang [Bangladesh, India, Indonesia, S Japan, S Korea, Laos, Malaysia,
Myanmar, Nepal, New Guinea, Sri Lanka, Vietnam].
One of us (Jeffrey) would treat “Gynostemma burmanicum,” G.
laxum, and G. longipes as synonyms of G. pentaphyllum var. pentaphyllum.

10b. Gynostemma pentaphyllum var. dasycarpum C. Y. Wu,
Acta Phytotax. Sin. 21: 362. 1983.
毛果绞股蓝 mao guo jiao gu lan
Gynostemma pentaphyllum f. dasycarpum (C. Y. Wu) W.
J. de Wilde & Duyfjes.
Fruit densely hispid and pubescent.
Forests; 1400–1700 m. S Yunnan [Indonesia (Borneo), Myanmar,
Thailand].

11. Gynostemma guangxiense X. X. Chen & D. H. Qin, Acta
Bot. Yunnan. 10: 495. 1988.
广西绞股蓝 guang xi jiao gu lan
Stems slender, glabrous, striate and sulcate. Tendrils filiform, apex bifid. Leaves alternate, pedate, with (3–)5–7 leaflets;
petiole 4–8 cm, glabrous; leaflet blade ovate-elliptic or obovate,
central leaflets 3–4.5 × 1.5–2.5 cm, lateral leaflets smaller,slightly oblique, membranous, glabrous or with sparse pubescence along midrib and lateral veins on both surfaces, base cuneate, margin serrate, apex shortly acuminate or obtuse; petiolule 3–5 mm. Plants monoecious. Male flowers greenish, ca. 2
m in diam., in axillary panicles 3–5 cm; peduncle filiform, 8–15mm, glabrous; bracteoles subulate, ca. 1.5 mm; pedicels fili-

CUCURBITACEAE

form, 2–3 mm; calyx segments deltoid, ca. 0.5 × 0.4 mm;
corolla segments lanceolate, 1–1.5 × ca. 0.5 mm; filaments very
short; anthers ovoid. Female flowers few, cymose; peduncle
slender, 6–14 mm, glabrous; pedicels 7–15 mm; calyx segments
lanceolate, ca. 1 × 0.3 mm; corolla segments lanceolate, ca. 2.5
× 0.5 mm, apex caudate-acuminate; ovary 2- or 3-locular, eachlocule with 1 ovule; styles 2 or 3, slightly connate at base; stigmas slightly bifid. Fruit greenish, obovate-triangular or 3angled-oblate, 4–5 × 5–6 mm, apex truncate, with persistentperianth and styles, glabrous; fruit stalk filiform, 7–18 mm.
Seeds 2 or 3, cream colored, oblong or subglobose, ca. 2.5 × 2.2mm, verruculose. Fl. Oct, fr. Nov–Dec.
● Forests on limestone mountains. Guangxi (Daxin, Longzhou).

12. Gynostemma compressum X. X. Chen & D. R. Liang,
Guihaia 11: 13. 1991.

Stem pubescent. Leaves papery, pedately 7–9-foliolate;petiole 4–8 cm; leaflets rhombic-elliptic or obovate-lanceolate,
median leaflets 5–12 × 2–4.5 cm, base attenuate, margin irregularly crenate, apex shortly acuminate; lateral leaflets smaller
than median leaflets, apex obtuse, abaxially hispid-pubescenton veins, adaxially sparsely pubescent; petiolule ca. 1 cm. Tendrils slender, 2-fid. Male flowers: panicle 10–20 cm, rachis andbranches pubescent; bracts linear, ca. 2 mm; pedicel filiform,ca. 4 mm; bracteole linear, ca. 0.7 mm; calyx segments ovate,ca. 1 × 0.5 mm, acute; corolla white; segments narrowly ovatelanceolate, ca. 2.5 × 1 mm, pubescent, 1-veined, apex long acuminate. Female flowers unknown. Fruit yellow-green, globose,
6–7 mm in diam., glabrous; fruiting pedicel filiform, (8–)15–20mm, glabrous. Seeds pale gray or brown, compressed cordate,ca. 3 × 3 mm, ca. 1 mm thick.
● Forests on valley sides; 1400–3200 m. Guangxi, Guizhou, S
Shaanxi, W Sichuan, NE and NW Yunnan.

扁果绞股蓝 bian guo jiao gu lan
Stems slender, sulcate, glabrous. Tendrils filiform, apex
usually bifid. Leaves pedate, with 7 leaflets; petiole 5–12 cm,
glabrous; leaflets abaxially gray-green, adaxially green, elliptic
or elliptic-ovate, membranous, base cuneate, margin crenulate, apex acuminate or acute, abaxially sparsely puberulent
along midrib and lateral veins, adaxially sparsely pilose, middle leaflets 3.6–6.5 × 2–2.25 cm, lateral leaflets smaller, outside oblique; petiolule 3–5 mm. Plants dioecious. Male flowers in panicles, axillary, 2–5 cm; peduncle filiform, 8–16 mm,
glabrous; bracteoles subulate, ca. 1.5 mm; pedicels 2–5 mm;flowers greenish, ca. 1.5 mm in diam.; calyx segments triangular, ca. 0.8 × 0.5 mm; filaments very short, connate; anthers
ovoid. Female flowers solitary or paired, axillary; peduncle filiform, 5–9 mm, glabrous; pedicels 5–10 mm; calyx segments ca.
1.3 × 0.4 mm; corolla segments linear-lanceolate, 2–3 × ca. 0.4mm, apex caudate-acuminate; ovary 2-locular, each locule with
1 ovule; styles 2, free; stigmas bifid. Fruit baccate, compressed,
obtriangular, 5–8 × 5–7 mm, glabrous, apex with persistentperianth and styles; fruit stalk filiform, 1.3–2.5 cm. Seeds
brownish, obtriangular, ca. 3 mm, apex ca. 3.5 mm in diam.,
both surfaces tuberculate, margin crenate and sulcate. Fl. Mar–
Apr, fr. Mar–May.
● Forests, moist limestone foothills; below 400 m. Guangxi(Longzhou).
This species is used in Chinese herbal medicine by the people of
Guangxi.

13. Gynostemma longipes C. Y. Wu, Acta Phytotax. Sin. 21:362. 1983.
长梗绞股蓝 chang geng jiao gu lan

15

One of us (Jeffrey) treats this taxon as a synonym of Gynostemma
pentaphyllum.

14. Gynostemma caulopterum S. Z. He, Acta Phytotax. Sin.
34: 207. 1996.
翅茎绞股蓝 chi jing jiao gu lan
Stem stout, 4–10 m high, 5–10 mm in diam., branched, 3angled, angles winged, wings 1.5–3 mm wide, glabrous orsparsely puberulent at nodes. Tendrils filiform, apex bifid.
Leaves pedate, 7-foliolate; petiole 5–8 cm, glabrous; leaflets
rhombic-elliptic or obovate-elliptic, papery, abaxially glabrous,
adaxially pilose, base cuneate, margin serrate, apex acuminate,mucronate; central leaflet 6–12 × 2–4 cm, lateral leaflets
smaller. Plants dioecious. Male flowers in axillary panicles, 7–9cm; bracteoles linear, ca. 1.5 mm; flowers ca. 2 mm in diam.;
pedicels 2–2.5 mm; calyx segments triangular, ca. 0.8 × 0.5mm; corolla segments ovate-lanceolate, ca. 1.5 × 1 mm; filaments very short, connate; anthers ovoid. Female flowers inaxillary or terminal panicles, 1.5–3.5 × 1.2–1.8 cm, glabrous;
pedicel 1–1.5 mm; basal bract 1, linear; calyx 5-lobed; segments triangular, ca. 0.5 × 0.4 mm; corolla 5-parted; segments
lanceolate, ca. 2 × 0.5 mm, apex acuminate; ovary compressed
globose, ca. 1.5 mm in diam., 2-locular, each locule with 1
ovule; styles 2, free; stigmas 3-fid. Berry greenish, compressed
globose, 2.5–3 × 5.5–6 mm, glabrous, with persistent corollaand styles; fruit stalks filiform, 3–6 mm. Seeds 2, creamcolored, broadly cordate, ca. 2 × 3 mm, spiniform-tuberculate.
Fl. and fr. Sep–Nov.
● Moist places along small stream valleys; 400–700 m. Guizhou(Renhuai, Ziyun).

4. NEOALSOMITRA Hutchinson, Ann. Bot. (Oxford), n.s., 6: 97. 1942.
棒锤瓜属 bang chui gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs or shrubs, climbing. Leaves simple or often 3–5-foliolate, leaflets sometimes 2-glandular at base. Tendrils simple or 2-fid.
Plants dioecious. Male flowers in paniculate or racemose inflorescences in axils of leaves; peduncle and pedicels filiform; calyx tube
cupular, 5-partite; segments oblong or oblong-lanceolate; corolla rotate, 5-partite; segments oblong; stamens 5, free; filaments short,CUCURBITACEAE

16

united at base; anthers oblong, 1-celled. Female flowers in lax axillary panicles or racemes; ovary 1-locular or imperfectly 3-locular;
ovules numerous, pendulous; styles 3, rarely 4; stigma semilunar. Fruit clavate or cylindric, terete or slightly 3-angled, apex broadly
truncate and 3-valved. Seeds imbricate, compressed, terminated by a very thin elongated wing, margins wavy-tuberculate, testa
crustaceous.
Eleven species: India to Polynesia and Australia; one species in China.

1. Neoalsomitra clavigera (Wallich) Hutchinson, Ann. Bot.
(Oxford), n.s., 6: 101. 1942.
棒锤瓜 bang chui gua
Zanonia clavigera Wallich, Pl. Asiat. Rar. 2: 28. 1831;
Alsomitra clavigera (Wallich) M. Roemer; A. integrifoliola(Cogniaux) Hayata; A. pubigera Prain; A. tonkinensis Gagnepain; Gynostemma elongatum Merrill; G. integrifoliolum Cogniaux; Hemsleya elongata (Merrill) Cogniaux; H. henryi Cogniaux; Neoalsomitra integrifoliola (Cogniaux) Hutchinson; N.
tonkinensis (Gagnepain) Hutchinson.
Herbs. Stem slender, much branched, pubescent or subglabrous. Leaf blade pedately 3–5-foliolate; petiole 1.5–2 cm, pubescent; leaflets abaxially pale green, adaxially green, oblongor oblong-lanceolate, median leaflet 7–14 × 3–7 cm, lateral
leaflets smaller, 1- or 2-glandular, membranous or thinly papery, both surfaces pubescent along veins, lateral veins 4 or 5
pairs, base obtuse, apex acuminate; petiolule 0.5–1 cm, densely
pubescent. Tendrils glabrous or pubescent, 2-fid. Male inflores-

cences paniculate, 12–30 cm, much branched; pedicel filiform,
4–8 mm, glabrous or pubescent; bracteole subulate-lanceolate,
1–2 mm, densely pubescent; calyx tube short; segments ovatelanceolate to lanceolate-linear, ca. 2 × 1 mm, hirsute-pubescent;corolla white; segments ovate, 2–4 × 1.5–3 mm, abaxiallydensely pubescent, apex acute; filaments ca. 0.8 mm; anthers
ovoid. Female flowers paniculate; calyx and corolla as in male
flowers; ovary subcylindric, ca. 10 mm, pubescent; styles 3;
stigma 2-lobed. Fruit capsular, cylindric, 4–8.5 × 1.5–2 cm, glabrous or pubescent, base obtuse, apex truncate, with numerous seeds. Seeds gray- or yellow-brown, substelliform, narrowly ovate, margin 4–7-dentate, 10–14 × 6–10 mm, both surfaces muriculate, apex with oblong, membranous, 15–22 × 6–8
mm wing. Fl. Sep–Nov, fr. Nov–Apr (following year).
Rain forests, broad-leaved forests, valleys, secondary forests.
Guangdong, Guangxi, Taiwan, SE Xizang (Mêdog), Yunnan [Bhutan,
Cambodia, NE India, Indonesia (Sumatra), Laos, Malaysia, Myanmar,
Nepal, Philippines, Thailand, Vietnam; Australia (Queensland), Pacificislands (Fiji, Solomon Islands)].

5. ZANONIA Linnaeus, Sp. Pl. 2: 1028. 1753.
翅子瓜属 chi zi gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Climbers, woody. Stem subglabrous. Leaves petiolate, undivided, margin entire. Tendrils simple or bifid. Plants dioecious; male
flowers in lax pendulous panicles; female flowers in racemes. Male flower: calyx tube shortly cupular; segments 3, rarely 4,
membranous, concave; corolla rotate; segments 5, apex narrow and inflexed in flower bud; stamens 5, free, all of same length,
inserted on a fleshy disk; filaments short, thick; anthers oblong, 1-celled; pistillode absent. Female flowers: calyx as in male flowers;
staminodes 5, very short, alternate with petals; ovary inferior, at first 3-loculed, ultimately 1-loculed; styles 3, spreading, apex bifid;
ovules in each locule 2 or many, pendulous, attached in 2 series to a fleshy parietal placenta. Fruit a capsule, large, narrowly
cylindric, clavate, 3-valved at truncate apex. Seeds pendulous, large, oblong, compressed, surrounded by a large membranous wing.
One species: Bhutan, Cambodia, China, India, Indonesia, Laos, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam.

1. Zanonia indica Linnaeus, Syst. Nat., ed. 10, 2: 1292. 1759.
翅子瓜 chi zi gua
Stem robust, branched, ± sulcate, glabrous. Leaves glabrous; leaf blade ovate-oblong, 8–16 × 5–10 cm, leathery, abaxially with distinct reticulate veins, lateral veins 3 or 4 pairs,
adaxially smooth, base rounded or slightly emarginate, margin
entire, apex acute; petiole 1.5–3 cm, glabrous. Male peduncle
slender; rachis ca. 16 cm, much branched; pedicel robust, 4–5mm, with articulation; calyx segments ovate-triangular, ca. 2
mm, glabrous, apex acute; corolla pale yellow-brown; segments
oblong, 3–3.5 × 1–1.5 mm, apex obtuse. Female peduncle 10–30 cm, sparsely 5–10-flowered; pedicels thick; calyx segments
ca. 4 mm, apex obtuse; corolla segments 6–8 mm; ovary obconic-cylindric, 10–12 mm. Fruit brownish, 6–10 × 2.5–5 cm,
finely granulate, base obtuse, apex truncate. Seeds oblong, ca. 2
× 1–1.5 cm; wing yellowish or white, surrounding central seed,
5–8 × 1.3–2 cm, base and apex rounded.

Riversides, mountain slopes; 300–900 m. W Guangxi, S Yunnan[Bhutan, Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam].

1a. Fruit 6–10 × 2.5–3 cm, glabrous; seed wing
5–6 × 1.3–1.5 cm .......................................... 1a. var. indica
1b. Fruit ca. 10 × 4.5–5 cm, densely
pubescent; seed wing ca. 8 × 2 cm ........ 1b. var. pubescens
1a. Zanonia indica var. indica
翅子瓜(原变种) chi zi gua (yuan bian zhong)Alsomitra simplicifolia Merrill; Juppia borneensis Merrill;
Zanonia indica subsp. orientalis W. J. de Wilde & Duyfjes.
Fruit 6–10 × 2.5–3 cm, glabrous. Seeds ca. 2 × 1 cm; wing
5–6 × 1.3–1.5 cm.
Riversides, mountain slopes; ca. 300 m. W Guangxi (Napo)[Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Philippines,
Sri Lanka, Thailand, Vietnam].

CUCURBITACEAE

171b. Zanonia indica var. pubescens Cogniaux in A. Candolle &C. Candolle, Monogr. Phan. 3: 927. 1881.

Fruit ca. 10 × 4.5–5 cm, densely pubescent. Seeds ca. 2 ×
1.5 cm; wing white, ca. 8 × 2 cm.

滇南翅子瓜 dian nan chi zi gua

Open forests on mountain slopes and dry valley sides; ca. 800 m.
S Yunnan (Mengla) [N India].

6. BOLBOSTEMMA Franquet, Bull. Mus. Natl. Hist. Nat., sér. 2, 2: 325. 1930.
假贝母属 jia bei mu shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, climbing, with a multi-tuberous base. Stems slender. Leaves cordate, 5-lobed, glandular at base. Tendrils 2-fid. Plants
dioecious; male flowers in panicle, female flowers often in panicle, sometimes solitary or fascicled. Male flowers: calyx rotate, 5partite; segments linear-lanceolate; corolla rotate, 5-partite; segments narrowly lanceolate, caudate-acuminate; stamens 5; filaments
free or connate at base in pairs; anthers subovoid, 1-celled; connective papillose on back. Female flowers: calyx and corolla as inmale flowers; ovary subglobose, verrucose, 3-locular; style short; stigmas 3, 2-lobed; ovules 2 in each locule, pendulous. Fruit narrowly ovoid, conical at apex, with or without spines, circumscissile-dehiscent above middle. Seeds 4–6, subovate, with membranous
wing at apex.
● Two species: disjunct between N China and SW China.

1a. Leaves palmately 5-fid, lobes 3–5-lobed; calyx segments and corolla segments ca. 2.5 mm; stamens free,connective not papillose; ovary rarely verrucose; fruit smooth ............................................................................... 1. B. paniculatum
1b. Leaves suborbicular; calyx segments and corolla segments to 6–8 mm; stamens connate at base offilaments in pairs, connective elongate; ovary densely verrucose; fruit spinescent ............................................ 2. B. biglandulosum
1. Bolbostemma paniculatum (Maximowicz) Franquet, Bull.
Mus. Natl. Hist. Nat., sér. 2, 2: 327. 1930.
假贝母 jia bei mu
Mitrosicyos paniculatus Maximowicz, Mém. Acad. Imp.
Sci. St.-Pétersbourg Divers Savans 9 [Prim. Fl. Amur.]: 113.
1859; Actinostemma multilobum Harms; A. paniculatum (Maximowicz) Cogniaux; Schizopepon fargesii Gagnepain.
Stem herbaceous, glabrous. Petiole slender, 1.5–3.5 cm;
leaf blade ovate-orbicular, 4–11 × 3–10 cm, palmately 5-partite;lateral lobes ovate-oblong, acute; central lobe oblong-lanceolate, acuminate; basal lobe with a gland at apex, glabrous or pilose on veins. Tendrils filiform, 2-fid. Inflorescence paniculate,
4–10 cm; pedicel filiform, 1.5–3.5 cm; calyx and corolla similar, yellow-green; segments ovate-lanceolate, ca. 2.5 cm; stamens free; filaments 0.3–0.5 mm; anthers ca. 0.5 mm; ovary
ovoid, 3-locular; ovules 2 in each locule. Fruit cylindric, 1.5–3
× 1–1.2 cm, circumscissile-dehiscent, conical at apex. Seeds 6,
8–10 × ca. 5 mm, testa rugose, margin denticulate, with a longmembranous wing at apex, wing 8–10 mm. Fl. Jun–Aug, fr.
Aug–Sep.
● Montane slopes. Gansu, Hebei, Henan, NW Hunan, Shaanxi,
Shandong, Shanxi, E and S Sichuan.
This plant is used medicinally.

2. Bolbostemma biglandulosum (Hemsley) Franquet, Bull.
Mus. Natl. Hist. Nat., sér. 2, 2: 328. 1930.
刺儿瓜 ci er gua
Branches slender, slightly pilose when young, ultimately
glabrous. Petiole slender, 2–5 cm; leaf blade broadly oblong or
suborbicular, 2.5–7 × 3–7 cm, membranous, glabrous, 5-lobed,
base broadly cordate, apex acute or obtuse, with a gland at apex
of lowest 2 lobes. Tendrils filiform, 2-fid, pilose or glabrous.
Male inflorescence paniculate, to 12 cm; rachis and pedicels
filiform; pedicels 1–1.5 cm; calyx segments linear-lanceolate;corolla segments pale yellow-green, narrowly lanceolate, 6–8 ×
0.8–1.2 mm, glabrous, apex long caudate; stamens inserted atbase of corolla tube, connate in pairs, 1 free; filaments ca. 0.8mm; anthers yellow, ca. 1 mm; connective caudate at apex, 1–
1.2 mm. Female flowers solitary; pedicels filiform, 10–15 mm;ovary densely verrucose. Fruit yellow-green, cylindric, 3.5–4
cm, covered with slender spines, spines 5–7 mm, circumscissile
when mature. Seeds irregularly ovate, margin irregularly dentate, ca. 10 × 7 mm, with membranous wing at apex, wing ca.
10 mm, apex lobate. Fl. Sep, fr. Oct.
● Forest margins, thickets under forests; 1000–1400 m. NW andSE Yunnan.

1a. Leaf blade suborbicular, ± as longas broad, 4–7 × 4–7 cm .................. 2a. var. biglandulosum
1b. Leaf blade broadly oblong, shorter
than broad, 2.5–3.5 × 3–4.5 cm.................................................... 2b. var. sinuatolobulatum
2a. Bolbostemma biglandulosum var. biglandulosum刺儿瓜(原变种) ci er gua (yuan bian zhong)Actinostemma biglandulosum Hemsley, Hooker’s Icon. Pl.
27: t. 2622. 1899; Hemsleya esquirolii H. Léveillé.
Leaf blade suborbicular, ± as long as broad, 4–7 × 4–7 cm.
Calyx segments linear-lanceolate; corolla segments narrowly
lanceolate, 6–8 × 1–1.2 mm.
● Forest margins; 1300–1400 m. SE Yunnan (Mengzi).

2b. Bolbostemma biglandulosum var. sinuatolobulatum C.
Y. Wu, Acta Phytotax. Sin. 6: 238. 1957.
波裂叶刺儿瓜 bo lie ye ci er gua
Leaf blade broadly oblong, shorter than broad, 2.5–3.5 ×
3–4.5 cm. Calyx- and corolla segments narrowly lanceolate, 6–
8 × 0.8–1 mm.
● Thickets under forests; ca. 1000 m. NW Yunnan (Lijiang).

CUCURBITACEAE

18

7. ACTINOSTEMMA Griffith, Account Bot. Coll. Cantor, 24. 1845.
盒子草属 he zi cao shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Mitrosicyos Maximowicz.
Herbs, scandent. Leaves petiolate; blade cordate-hastate, cordate-ovate, broadly ovate, or lanceolate-triangular, unlobed or 3–5lobed, margins dentate. Tendrils 2-fid or rarely simple. Plants monoecious or rarely bisexual. Flowers unisexual. Male inflorescence
racemose or paniculate, rarely flowers solitary or paired; calyx rotate, segments linear-lanceolate; corolla rotate, segments lanceolate,acuminate; stamens 5(or 6), free; filaments short, filiform; anthers subovoid, extrorse, basifixed, 1-celled; anther cell oblong; connective papillose on back. Female flowers solitary or fasciculate; calyx and corolla as in male flowers; ovary ovoid, verrucose, 1locular; style short; stigmas 3, reniform; ovules 2(–4), attached to wall near apex of locule, pendulous. Fruit ovoid, conical at apex,
semi-superior, echinate, circumscissile-dehiscent near middle. Seeds 2–4, compressed, testa rugose, margin denticulate.
One species: E Asia and India.

1. Actinostemma tenerum Griffith, Account Bot. Coll. Cantor,
25. 1845.
盒子草 he zi cao
Branches slender, slightly pilose when young, ultimately
glabrous. Petiole filiform, slightly puberulent, 2–6 cm; blade
very variable, ovate or sagittate-ovate, 3–12 × 2–8 cm, abaxially slightly pilose, ultimately glabrous, adaxially indistinctly
punctate-scabrid, 3-lobed, base deeply emarginate, margin minutely denticulate or coarsely serrate, apex long, acute-acuminate. Tendrils filiform, 2-fid. Male inflorescence racemose,
sometimes paniculate, shortly villous, many flowered, mainrachis 1–13 cm, shortly villous; bract linear, ca. 3 mm, villous;calyx segments linear-lanceolate, 2–3 × 0.5–1 mm, margindentate; corolla segments lanceolate, 2–3 × 1–1.5 mm at base,scarcely villous, 1- or 3-veined, apex narrowly subulate; staminal filaments slightly dilated at apex, ca. 0.5 mm, pilose; anthers ca. 0.3 mm. Female flowers solitary or fasciculate; calyx
and corolla as in male flowers; ovary ovoid, verrucose. Fruitovoid or oblong-ellipsoid, 1.6–2.5 × 1–2 cm, verrucose, operculate, 2–4-seeded. Seed plano-convex, broadly ovoid, 11–13 ×
8–9 mm, 3–4 mm thick, both surfaces reticulate, base slightly
attenuate. Fl. Jul–Sep, fr. Sep–Nov.
Growing on watersides. Anhui, Fujian, Guangxi, Hebei, Henan,
Hubei, Hunan, Jiangsu, Jiangxi, Liaoning, Shandong, Sichuan, S Xizang, W Yunnan, Zhejiang [India, Japan, Korea, Laos, Thailand, Vietnam].

The whole plant and its seeds are used medicinally.

1a. Flowers unisexual; stamens 5 .................... 1a. var. tenerum
1b. Flowers bisexual; stamens 6 ............... 1b. var. yunnanense
1a. Actinostemma tenerum var. tenerum盒子草(原变种) he zi cao (yuan bian zhong)Actinostemma japonicum (Miquel) Miquel; A. lobatum(Maximowicz) Franchet & Savatier; A. lobatum var. japonicum
Maximowicz; A. lobatum f. longilobum Komarov; A. lobatum
var. palmatum Makino; A. lobatum f. subintegrum Komarov; A.
palmatum (Makino) Makino; A. parvifolium Cogniaux; A. racemosum (Maximowicz) Cogniaux; Mitrosicyos lobatus Maximowicz; M. racemosus Maximowicz; Pomasterion japonicum
Miquel.
Flowers unisexual. Stamens 5.
Grasslands at watersides. Anhui, Fujian, Guangxi, Hebei, Henan,
Hubei, Hunan, Jiangsu, Jiangxi, Liaoning, Shandong, S Xizang, W
Yunnan, Zhejiang [India, Japan, Korea, Laos, Thailand, Vietnam].

1b. Actinostemma tenerum var. yunnanense A. M. Lu & Zhi
Y. Zhang, Bull. Bot. Res., Harbin 4(2): 127. 1984 [“yunnanensis”].
云南盒子草 yun nan he zi cao
Flowers bisexual. Stamens 6.
● W Yunnan.

8. INDOFEVILLEA Chatterjee, Nature 158: 345. 1946.
藏瓜属 zang gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Woody climbers, large. Stem angular-sulcate. Leaves petiolate; leaf blade ovate, cordate, entire, pedately veined at base. Tendrils 2-fid. Plants dioecious. Male flowers in panicle; rachis elongate; pedicels short; bracts linear, apex acuminate; calyx segments 5,
3- or 4-veined, acute; corolla segments 5, shorter than calyx segments, 3–5-veined; stamens 5, 4 in pairs, 1 free; filaments very short;anthers reniform, 1-celled, pubescent; staminodes and rudimentary ovary absent. Fruit oblong, indehiscent, in clusters of 3–6 on avery long peduncle, with a thick woody rind. Seeds many, large, ovate, compressed, faintly margined.
One species: SW China, NE India.

1. Indofevillea khasiana Chatterjee, Nature 158: 345. 1946.
藏瓜 zang gua

Stem branched, robust, pubescent at first, ultimately glabrous. Tendrils smooth, glabrous, to 20–30 cm. Petiole robust,
1.5–8 cm, pubescent at first, later glabrous; leaf blade broadly

CUCURBITACEAE

ovate-cordate, 15–25 × 12–20 cm, leathery, margin entire or
rarely in younger leaves distinctly and minutely toothed nearbase, scabrous on veins, otherwise glabrous on both surfaces,
apex shortly acuminate. Male panicle to 17 cm; rachis andbranches scabrous; bracts 5–9 mm; bracteoles 2–3 mm; calyx
segments ovate-lanceolate, longer than corolla segments, ca. 6

19mm; corolla segments ovate-lanceolate, ca. 4 mm, outsidesparsely and minutely hairy, inside papillose. Fruit oblong, 3–6
in clusters on a long stalk, ca. 30 cm, indehiscent with a thickwoody rind. Seeds 3.5–4 × ca. 2 cm, ca. 0.5 cm thick. Fl. and fr.
Aug.
Open forests on mountain slopes; ca. 900 m. S Xizang [NE India].

9. SIRAITIA Merrill, Pap. Michigan Acad. Sci. 19: 200. 1934.
罗汉果属 luo han guo shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Neoluffa Chakravarty.
Herbs, perennial, scandent, verrucose-glandular. Tuber spheroid, enlarged. Stems angular-sulcate. Leaf blade unlobed, margin
sparsely dentate, abaxially pilose or densely tomentose. Tendrils spiralled above and below branch, 2-fid. Plants dioecious. Male
flowers: inflorescence paniculate, bracteate; calyx segments 5, triangular, densely pubescent; corolla segments 5, lanceolate orobovate-lanceolate, base often with 5 scales; stamens 5, 4 connected in pairs, 1 free; anthers sigmoid. Female flowers solitary, paired,or 3 or 4 in raceme; staminodes 5, glanduliform; ovary ovoid; ovules numerous, horizontal; style stout, apex 3-lobed; stigma 2-lobed.
Fruit subglobose or cylindric. Seeds subovoid or ovoid, winged, wing corky. 2n = 28.
Four species: China, India (Sikkim), Indonesia, Malaysia, Thailand, Vietnam; three species (one endemic) in China.

1a. Male inflorescence cymose-paniculate, many flowered ............................................................................................ 1. S. sikkimensis
1b. Male inflorescence racemose or paniculate, 6–10-flowered.
2a. Seeds with 3-layered wing, wing marginally obtusely dentate; calyx segments 3–5 × 7–9 mm .......................... 3. S. siamensis
2b. Seeds with 2-layered wing, wing marginally only sparsely dentate; calyx segments 3–4 × 3–4 mm ............... 2. S. grosvenorii
1. Siraitia sikkimensis (Chakravarty) C. Jeffrey, Kew Bull. 36:
737. 1982.
锡金罗汉果 xi jin luo han guo
Neoluffa sikkimensis Chakravarty, J. Bombay Nat. Hist.
Soc. 50: 895. 1952.
Stem angular, tomentose. Petiole 5–8 cm, tomentose; leaf
blade cordate, 7–15 × 7–15 cm, abaxially tomentose, adaxiallyglabrous, margin entire, lowest lateral veins running parallelwith margin of basal lobes. Male flowers yellow, 20–35
crowded on 8–14 cm axillary peduncle; rachis 3–5 cm; pedicels
1–2 cm; bracts leaflike, 7–15 mm; calyx tube campanulate, 10–
15 mm in diam., tomentose; segments acute, 3–5 mm; corollasegments oblong-ovate, 8–10 × 3–4 mm, entire, ± constricted atapex, apex acute; stamens free, inserted at base of calyx tube,
1–2 mm; filaments 0.5–1 mm; anthers 1-celled, sinuate; connective broad, granular. Female flowers and fruit unknown.
S Yunnan (Mengla) [India (Sikkim)].

scales; segments triangular, ca. 4.5 × 3 mm, 3-veined, apex longacuminate; corolla yellow; segments oblong, 10–15 × 7–8 mm,
5-veined, apex acute; filaments puberulent, ca. 4 mm; anthersca. 3 mm. Female flowers solitary or 2–5 on 6–8 mm peduncle; calyx and corolla as in male flowers but slightly larger;
staminodes 2–2.5 mm; ovary oblong, 10–12 × 5–6 mm, densely
yellow-brown velvety, base obtuse-rounded; style ca. 2.5 mm;stigmas 3, enlarged, ca. 1.5 mm. Fruit globose or oblong, 6–11
× 4–8 cm, densely yellow-brown velvety and black glandularscaly, ultimately glabrous. Seeds numerous, pale yellow,
broadly ovate, compressed, 15–18 × 10–12 mm, base obtuserounded, with 2-layered wings, wings sinuate. Fl. May–Jul, fl.
Jul–Sep.
● Forests on mountain slopes, riversides, thickets. Guangdong,
Guangxi, Guizhou, S Hunan, Jiangxi.
The fruit is used medicinally.

3. Siraitia siamensis (Craib) C. Jeffrey ex S. Q. Zhong & D.
Fang, Guihaia 4: 23. 1984.

2. Siraitia grosvenorii (Swingle) C. Jeffrey ex A. M. Lu & Zhi
Y. Zhang, Guihaia 4: 29. 1984.

翅子罗汉果 chi zi luo han guo

罗汉果 luo han guo

Thladiantha siamensis Craib, Bull. Misc. Inform. Kew
1914: 7. 1914; Momordica tonkinensis Gagnepain.

Momordica grosvenorii Swingle, J. Arnold Arbor. 22: 198.
1941; Thladiantha grosvenorii (Swingle) C. Jeffrey.
Plant: all parts with yellow-brown pubescence and black
glandular scales. Root enlarged, fusiform or subglobose. Stemand branches slightly robust. Petiole 3–10 cm; leaf blade ovatecordate, 12–23 × 5–17 cm, membranous, apex acuminate orlong acuminate; sinus semicircular or broadly ovate-cordate.
Male flowers: inflorescence racemose, 6–10-flowered; peduncle 7–13 cm; pedicels slender, 5–15 mm; calyx tube broadly
campanulate, 4–5 × ca. 8 mm, usually with 3 membranous

Plants to 20 m; all parts densely yellow-brown pubescentand red (black when dry) glandular-scaly. Roots enlarged. Petiole 3.5–10 cm; leaf blade ovate-cordate, 10–27 × 2–5.5 cm,
membranous, palmately 5–7-veined. Male flowers: 5–15(tomany) in a raceme or panicle; peduncle 2–12 cm; pedicels 15–
30 mm; calyx tube broadly campanulate, 12–15 mm in diam.;segments ovate-triangular, 3–5 × 7–9 mm, 3-veined, apex obtuse; corolla pale yellow, 3.4–4 cm in diam.; segments ovateoblong, 15–20 × 9–13 mm, margin with glandular hairs, basewith 3 membranous scales, 5-veined, apex obtuse; filaments

CUCURBITACEAE

20

sparsely glandular hairy. Female flowers solitary or paired;calyx and corolla as in male flowers but smaller; staminodes(3–)5; corolla base with 3 scales; ovary ovoid, 12–15 × 9–10mm, densely velvety and glandular-scaly, base obtuse-rounded,
apex truncate; style 4–5 mm, glabrous, 3-lobed; stigma reniform. Fruit subglobose, ca. 6 cm in diam., velvety at first, glabrescent. Seeds numerous, pale brown, ovoid, 12–14 × 11–13mm, ca. 4 mm thick, with 3-layered wings; wing corky, margin
irregularly dentate, median 3–5 mm wide, lateral 1–2 mm wide.
Fl. Apr–Jun, fr. Jul–Sep.
Forests on mountain slopes; 300–700 m. W Guangxi, SE Yunnan[Indonesia, Malaysia, Thailand, N Vietnam].
The tuber is used medicinally.

10. SINOBAIJIANIA C. Jeffrey & W. J. de Wilde, Bot. Zhurn. 91: 769. 2006.
白兼果属 bai jian guo shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, perennial, scandent. Root spheroid, enlarged. Stem and branches sulcate and angular, without verrucose glands, sparsely
puberulent. Leaf blade ovate-cordate or triangular-cordate, membranous, abaxially hispid on veins, adaxially strigose-punctate, margin entire or 2- or 3-lobed. Male inflorescence racemose, 5–8-flowered, bracteate; calyx segments triangular-ovate, valvate, sparsely
pilose; corolla oblong or ovate; stamens 5; anthers curved. Female flowers 1–3, ovoid or cylindric; ovules numerous, horizontal.
Fruit subglobose or cylindric. Seeds ovate or ovate-oblong, apex truncate or obtuse, unwinged. 2n = 32.
Four species: S China, Laos, Thailand; three species (two endemic) in China.
One of us (Lu) does not recognize “micro-genera” such as this; however, for the purposes of this Flora, the treatment by the co-author (Jeffrey)
is followed.

1a. Leaves 3–5-parted; fruit subglobose .......................................................................................................................... 3. S. taiwaniana
1b. Leaves entire, undulate, crenate, or lobed; fruit cylindric.
2a. Leaves entire, undulate; seeds triangular-ovate, apex obtuse and apiculate .......................................................... 1. S. decipiens
2b. Leaves crenate or lobed; seeds ovate, apex broadly truncate ........................................................................... 2. S. yunnanensis
1. Sinobaijiania decipiens C. Jeffrey & W. J. de Wilde, Bot.
Zhurn. 91: 769. 2006.

2. Sinobaijiania yunnanensis (A. M. Lu & Zhi Y. Zhang) C.
Jeffrey & W. J. de Wilde, Bot. Zhurn. 91: 769. 2006.

白兼果 bai jian guo

云南白兼果 yun nan bai jian guo

Stem and branches setose and pubescent at first, ultimately
glabrous. Petiole 2–6 cm; leaf blade narrowly ovate-cordate, 8–
15 × 5–10 cm, membranous or papery, abaxially sparsely puberulent or subglabrous, adaxially setose at first, ultimately strigose-punctate, pubescent on veins, base cordate, margin entire
or callose-serrate, apex acuminate; sinus semicircular or suborbicular, 1.5–2.5 cm deep, 2–3 cm wide. Tendrils 2-fid. Plants
dioecious. Male inflorescence racemose or paniculate; rachis 3–
14 cm, pubescent, 7–15-flowered; pedicel 5–10 mm, pubescent,base with a lanceolate or narrowly ovate, 2–3 mm bract; calyx
tube cupular, 2.5–3 mm, ± puberulent; segments lanceolate, 6–
7.5 mm, base 2–3 mm wide, 3-veined, apex acuminate; corolla
yellow; segments narrowly ovate, 6–6.5 mm, apex acute; stamens 4 in pairs and 1 free; filaments ca. 1 mm; anthers terete,
1-celled, ca. 3 mm, cells curved, puberulent. Female flowers
solitary, or 2- or 3-flowered at apex of peduncle; pedicel ca. 5mm, puberulent; calyx and corolla as in male flowers; ovary
oblong, ca. 8 × 4 mm, densely hispid or pubescent; style ± long,
filiform, ca. 7 mm, 3-fid; stigma 2-lobed, enlarged. Fruiting
pedicel stout, sulcate-angular, 2–2.5 cm, densely pubescent andsetose, ultimately glabrous; fruit red-brown when mature, oblong or subglobose, 5.5–6.5 × 4–5 cm, pubescent at first, ultimately glabrous. Seeds numerous, pale yellow-brown, triangular-ovate, ca. 8 × 7 cm, ca. 3 mm thick, both surfaces smooth,base obtuse-rounded, constricted above, apex truncate. Fl. Jun–
Aug, fr. Sep–Dec.

Siraitia borneensis (Merrill) C. Jeffrey ex A. M. Lu & Zhi
Y. Zhang var. yunnanensis A. M. Lu & Zhi Y. Zhang, Guihaia
4: 31. 1984; Baijiania yunnanensis (A. M. Lu & Zhi Y. Zhang)
A. M. Lu & J. Q. Li; S. borneensis var. lobophylla A. M. Lu &Zhi Y. Zhang.

● Forests, river valleys. Guangdong, SE Xizang (Mêdog), SE
Yunnan (Xichou).

Tuber complanate-spheroid, 10–13 × 6–8 cm. Stem angular-sulcate, sparsely puberulent. Petiole 3–9 cm; leaf blade
ovate-cordate, 5–13 × 4–10 cm, membranous, abaxially pilose,
hispid on veins, adaxially scabrous, margin irregularly dentate
or slightly 3-lobed, apex acute. Tendrils 2-fid. Male flowers: 8
in raceme; peduncle 4–6 cm; bracts broadly obovate, 5–6 × 3–4mm, pubescent; calyx segments triangular-ovate or ovate, 5.5–8
× 3–5 mm, 3-veined, apex acuminate; corolla segments oblong,
10–13 × 7–8 mm, villous, 5-veined, base truncate, apex obtuse;
filaments 3–3.5 mm; anthers curved, 3.5–5 mm. Female flowers
solitary or paired; pedicel 1–1.5 cm, pubescent; ovary cylindric,ca. 10 × 4 mm, densely villous. Fruit cylindric, ca. 5 × 3.3 cm.
Seeds numerous, ovoid or ovoid-oblong, 7–8.5 × 4.5–6 mm, ca.
1.5 mm thick.
Valley sides, forests; 1000–1800 m. S and SW Yunnan [N Laos].
De Wilde and Duyfjes (Blumea 51: 497. 2006) cited evidence
suggesting that the type of Siraitia borneensis var. lobophylla may
represent a species of Sinobaijiania distinct from S. yunnanensis.

3. Sinobaijiania taiwaniana (Hayata) C. Jeffrey & W. J. de
Wilde, Bot. Zhurn. 91: 770. 2006.
台湾白兼果 tai wan bai jian guo

CUCURBITACEAE

Thladiantha taiwaniana Hayata, J. Coll. Sci. Imp. Univ.
Tokyo 30(1): 119. 1911; Baijiania taiwaniana (Hayata) A. M.
Lu & J. Q. Li; Siraitia taiwaniana (Hayata) C. Jeffrey ex A. M.
Lu & Zhi Y. Zhang.
Stem and branches sulcate-angular, subglabrous. Petiole
ca. 3 cm; leaf blade oblong-cordate or ovate-cordate, 5–12 × 5–
10 cm, abaxially sparsely setose at first, ultimately glabrous,
adaxially scabrous, punctate, 5-veined, base deeply cordate,21

margin irregularly dentate, 3-lobed, apex acute; median lobe
oblong, ca. 6 × 3.5 cm. Tendrils simple. Male flowers: inflorescence racemose, axillary, 6–7 cm; bracts broadly obovate, ca.
7 × 5 mm, pubescent; calyx tube ca. 4 mm in diam., pubescent;segments 5, triangular-ovate, ca. 4 × 2.5 mm, apex acuminate;
corolla segments 5, oblong, apex obtuse. Fruit globose, ca. 2.5
cm in diam., pubescent.
● Taiwan.

11. THLADIANTHA Bunge, Enum. Pl. Chin. Bor. 29. 1833.
赤瓟属 chi bao shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, perennial, climbing, scandent, or prostrate. Roots usually tuberous. Stems herbaceous, scabrous. Tendrils simple or 2-fid.
Leaves mostly simple, vary rarely pedately 3–7-foliolate; leaf blade cordate, margin dentate. Plants dioecious. Male flowers incymes-racemes or rarely solitary; calyx tube broadly campanulate or cupular; segments 5, linear, lanceolate, ovate-lanceolate, oroblong, 1–3-veined; corolla yellow, campanulate, 5-partite; segments oblong, broadly ovate, or obovate, usually 5–7-veined, entire;stamens 5, inserted on calyx tube, free, often 4 in pairs, other 1 free; filaments short; anthers oblong or ovoid, all 1-celled; anther cellerect; rudimentary ovary glandlike. Female flowers solitary or 2- or 3-fascicled on a short peduncle; calyx and corolla as in male
flowers; ovary ovoid, oblong, or fusiform, smooth or verrucose; ovules numerous, horizontal; style 3-fid; stigmas 3, 2-lobed, reniform. Fruit fleshy, indehiscent, smooth or verrucose, many ribbed or not ribbed. Seeds numerous, horizontal.
Twenty-three species: Bhutan, China, India, Indonesia, Japan (introduced), Korea, Laos, Myanmar, Nepal, Thailand, Vietnam; 23 species (19endemic) in China.

1a. Bracts in male flowers flabelliform, dentate.
2a. Calyx segments entire, 0.6–3 mm wide ................................................................................................................. 1. T. cordifolia
2b. Calyx segments dentate or irregularly lobed, (3–)4–11 mm wide.
3a. Calyx segments (3–)4–6 mm wide, with 1–3 triangular lobes .................................................................. 2. T. grandisepala
3b. Calyx segments to 11 mm wide, flabelliform, margin irregularly lobed .................................................... 3. T. medogensis
1b. Male flowers without flabelliform bracts.
4a. Ovary and fruit scaly or verrucose.
5a. Leaf blade ovate-lanceolate or narrowly ovate-triangular; ovary and fruit verrucose .................................. 23. T. longifolia
5b. Leaf blade ovate-cordate; fruit with thick scales.
6a. Ovary and fruit ovoid-globose, apex with a long rostrum, scales at base of fruit less than 5 mm,
not adnate to pedicel ................................................................................................................................ 21. T. pustulata
6b. Ovary and fruit ovoid-oblong, apex without a rostrum, scales at base of fruit to 10 mm,
adnate to pedicel ........................................................................................................................................... 22. T. henryi
4b. Ovary and fruit not scaly or verrucose.
7a. Leaves palmately deeply 5-lobed .......................................................................................................... 20. T. palmatipartita
7b. Leaves not palmately deeply 5-lobed.
8a. Male flowers congested at apex of rachis into pseudocapitulum.
9a. Calyx segments lanceolate, apex acute, 1-veined .............................................................................. 18. T. capitata
9b. Calyx segments linear, apex obtuse, 3-veined ..................................................................................... 19. T. davidii
8b. Male flowers in cyme-panicle, panicle, raceme, or rarely solitary, but not capitellate.
10a. Plants sparsely hairy or subglabrous.
11a. Calyx segments 1-veined.
12a. Basal lateral vein along base of leaf blade; calyx segments linear, over 9 mm .............. 13. T. oliveri
12b. Basal lateral vein not along base of leaf blade; calyx segments triangularlanceolate, 4–5 mm ....................................................................................................... 14. T. montana
11b. Calyx segments 3-veined.
13a. Tendrils 2-fid, leaves not lobed or lobed ........................................................................ 17. T. dentata
13b. Tendrils simple, leaves not lobed.
14a. Leaf blade broadly ovate-cordate; tendrils filiform; pedicel in male flowers
1.2–2.5 cm; fruit with a long beak ..................................................................... 15. T. maculata
14b. Leaf blade narrowly ovate or narrowly ovate-lanceolate; tendrils robust;
pedicel 0.5–1 cm; fruit without beak .................................................................. 16. T. punctata
10b. Plants with dense multicellular setae, very rarely subglabrous.

CUCURBITACEAE

2215a. Leaves simple, unlobed or margin lobed or serrate, or pedately 3–5-foliolate ........................ 4. T. hookeri
15b. Leaves unlobed, rarely irregularly lobed.
16a. Tendrils simple.
17a. Male flowers solitary or in a short raceme, bracts leaflike ........................................ 5. T. dubia
17b. Male flowers without bracts.
18a. Petiole less than 1.5 cm, leaf blade lanceolate or narrowly ovate ........... 6. T. sessilifolia
18b. Petiole over 2 cm, leaf blade ovate-cordate ................................................ 7. T. villosula
16b. Tendrils 2-fid.
19a. Male flowers dimorphic, solitary, and in raceme with (2 or)3(–5) flowers
in axil of same leaf ....................................................................................... 8. T. dimorphantha
19b. Male flowers in panicle or raceme.
20a. Stem and leaves with pale yellow multicellular setae or very rarely
subglabrous, sinus at base of leaf closed; fruit orange-red whenmature .......................................................................................................... 9. T. nudiflora
20b. Stem and leaves with gray-white multicellular setae, sinus at baseof leaf open; fruit whitish yellow when mature.
21a. Calyx segments broadly lanceolate or obovate-oblong,
10–15 mm .................................................................................... 10. T. longisepala
21b. Calyx segments triangular-lanceolate, less than 10 mm.
22a. Ovary and fruit with dense yellow-brown spiniform setae;
seeds 6.5–7 mm, sparsely irregularly verrucose on surface ... 11. T. setispina
22b. Ovary and fruit with long setae; seeds 4.5–5 mm, notverrucose on surface ............................................................ 12. T. lijiangensis
1. Thladiantha cordifolia (Blume) Cogniaux in A. Candolle &C. Candolle, Monogr. Phan. 3: 424. 1881.

Streamsides, forests; 800–2600 m. Guangdong, Guangxi, Sichuan,
Yunnan [India, Indonesia, Laos, Myanmar, Nepal, Thailand, Vietnam].

大苞赤瓟 da bao chi bao

De Wilde and Duyfjes (Blumea 51: 513–514. 2006) considered
Thladiantha calcarata var. tonkinensis (as T. tonkinensis Gagnepain)
and T. globicarpa to be species distinct from T. cordifolia, differing intheir smaller, non-fenestrate fruit and the latter from the former by its
narrower calyx segments.

Luffa cordifolia Blume, Bijdr. 929. 1826; Gymnopetalum
piperifolium Miquel; Thladiantha calcarata C. B. Clarke; T.
calcarata var. subglabra Cogniaux; T. calcarata var. tonkinensis Cogniaux; T. cordifolia var. tomentosa A. M. Lu & Zhi Y.
Zhang; T. cordifolia var. tonkinensis (Cogniaux) A. M. Lu &Zhi Y. Zhang; T. globicarpa A. M. Lu & Zhi Y. Zhang; T.
omeiensis T. Y. Zhu; T. tonkinensis Gagnepain; Trichosanthes
javanica Miquel.
Vines, climbing, herbaceous, pubescent. Stems muchbranched, robust, angular-sulcate. Petiole slender, 4–10(–12)
cm; leaf blade ovate-cordate, 8–15 × 6–11 cm, membranous or
papery, adaxially scabrous, pubescent, or with setae on an enlarged base, veins densely pubescent, both surfaces densely
pale yellow pubescent, basal vein along margin of base of
blade, base cordate, margin irregularly dentate, apex acuminate
or shortly so; sinus at base often open, sometimes closed, 1–3
cm deep, 0.5–2 cm wide. Tendrils simple, pubescent at first,
glabrescent. Male flowers: 3 to several in dense short raceme;
peduncle robust, 4–15 cm, pubescent; bracts imbricate, flabelliform, 1.5–2 cm, both surfaces pubescent; pedicels slender, ca.
5 mm; calyx tube campanulate, 5–6 mm, 5-partite; segments
linear, ca. 10 × 1 mm, 1-veined, apex caudate-acuminate; corolla segments ovate or elliptic, ca. 17 × 7 mm, apex shortly
acuminate or acute; filaments ca. 4 mm; rudimentary ovary
semiglobose. Female flowers solitary; calyx and corolla as inmale flowers; ovary oblong, base obtuse, pubescent; styles 3fid; stigmas enlarged, reniform, 2-lobed. Fruit oblong, 3–5 × 2–
3 cm, with scabrous rind, slightly pubescent, with 10 longitudinal striae, obtuse at both ends. Seeds broadly ovate, 4–5 ×
3–3.5 mm, ca. 2 mm thick, reticulate. Fl. and fr. May–Nov.

2. Thladiantha grandisepala A. M. Lu & Zhi Y. Zhang, Bull.
Bot. Res., Harbin 1(1–2): 67. 1981.
大萼赤瓟 da e chi bao
Vines, climbing, herbaceous. Stems and branches slender,
angular-sulcate, at first densely pubescent, glabrescent. Petiole
4–8 cm; leaf blade narrowly ovate-cordate, 10–16 × 6–11 cm,
membranous, adaxially scabrous, with setae on an enlarged
base, veins densely pubescent, basal vein along margin of baseof leaf, base cordate, margin dentate, apex shortly acuminate;sinus at base open, semicircular, 1–2 cm deep, 0.5–1 cm wide.
Male flowers: 5–9 in pedunculate raceme; bracts flabelliform,
12–15 × 15–17 mm, both surfaces pubescent, margin irregularly dentate; pedicels slender, 5–10 mm, pubescent; calyx tube
campanulate, ca. 4 mm; segments broadly lanceolate to narrowly ovate, slightly irregular, ca. 1.5 cm × 4–6 mm, distinctly
3-veined, apex usually irregularly 2- or 3-lobed; corolla segments narrowly oblong, ca. 2 × 1 cm, apex shortly acuminate;filaments pubescent; rudimentary ovary semiglobose. Female
flowers solitary; pedicels 2–5 cm, pubescent; calyx segments
broadly lanceolate or narrowly ovate, to 18 × ca. 6 mm, 3veined, margin often 2- or 3-lobed, apex acuminate; corollasegments ca. 3 × 1.4 cm; ovary narrowly oblong, 15–18 × 4–6
mm, densely yellow villous, base narrow, apex subtruncate;style filiform, 3-fid; stigma enlarged, 2-lobed. Fruiting pedicelrobust, 3–5 cm, subglabrous; fruit elliptic, 2.5–3 × ca. 1.5 cm,
pubescent, obtuse at both ends. Fl. Jun–Aug, fr. Aug–Oct.
● Mountain slopes, forests; 2100–2400 m. Yunnan.

CUCURBITACEAE

3. Thladiantha medogensis A. M. Lu & J. Q. Li, Acta Bot.
Yunnan. 14: 133. 1992.
墨脱赤瓟 mo tuo chi bao
Herbs, scandent. Stem slender, angular-sulcate, sparsely
puberulent. Petiole 3.5–5 cm, pubescent and sparsely setose;
leaf blade ovate-cordate, 8–11 × 5–5.6 cm, papery or submembranous, abaxially sparsely pubescent and setose, adaxiallydensely setose at first, ultimately punctate-scabrous, basal lateral veins along margin of sinus, base emarginate, margin callose-dentate, apex shortly acuminate. Tendrils simple, pubescent. Male flowers in a raceme; peduncle robust, 8–24 cm;
bracts ca. 2 × 2.2 cm, margin irregularly 5- or 6-obtusely dentate; pedicel slender, 7–10 mm, sparsely pubescent; calyx tube
campanulate, ca. 2.5 × 3.5 mm; segments ca. 10 × 11 mm, 3veined, margin irregularly 3–5-dentate; corolla segments narrowly oblong, 8–9 × 3–3.5 mm, apex acuminate; filaments ca.
1.5 mm, sparsely pubescent; anthers oblong, ca. 3 mm; rudimentary ovary subglobose, ca. 2 mm in diam. Fruit solitary,subcylindric, ca. 3.2 × 2 cm, longitudinally striate, rounded atapex and base.
● About 1800 m. SE Xizang (Mêdog).

4. Thladiantha hookeri C. B. Clarke in J. D. Hooker, Fl. Brit.
India 2: 631. 1879.
异叶赤瓟 yi ye chi bao
Hemsleya tonkinensis Cogniaux; H. trifoliolata Cogniaux;
H. yunnanensis Cogniaux; Thladiantha digitata H. Léveillé; T.
heptadactyla Cogniaux; T. hookeri var. heptadactyla (Cogniaux) A. M. Lu & Zhi Y. Zhang; T. hookeri var. palmatifolia
Chakravarty; T. hookeri var. pentadactyla (Cogniaux) A. M. Lu& Zhi Y. Zhang; T. hookeri f. quinquefoliata Chakravarty; T.
hookeri f. trifoliolata (Cogniaux) Chakravarty; T. pentadactyla
Cogniaux; T. trifoliolata (Cogniaux) Merrill.
Vines, climbing, herbaceous. Tuber oblate-spheroid. Stems
2–4(–10) m, to 10 mm in diam., much branched, subglabrous.
Petiole 3–6 cm, glabrous or puberulent; leaf blade pedately 3–
7-foliolate or undivided; leaflets oblong-lanceolate, broadly lanceolate, lanceolate, or linear-lanceolate, median leaflet 6–15 ×
1–3 cm, membranous, abaxially subglabrous, adaxially scabrous, margin dentate, apex acuminate. Tendrils filiform, simple,
smooth. Male flowers in raceme or solitary; rachis filiform, 2–4
cm, glabrous, ebracteate; pedicel of solitary flowers to 2.5 cm;
calyx tube broadly campanulate, 3–4 × 6–7 mm, sparsely pubescent; segments erect, narrowly triangular, ca. 4 × 1.5 mm, 3veined; corolla segments ovate, 10–12 × ca. 5 mm; filaments 2–
3 mm, glabrous; anthers oblong, ca. 2 mm. Female flowers
solitary; pedicel filiform, 2–4 cm, at first puberulent, ultimately
glabrous; calyx and corolla as in male flowers; calyx segments
ca. 10 mm; corolla segments ca. 2 cm; ovary fusiform, 1–2 cm
× 2–3 mm, densely yellow-brown pubescent, attenuate at both
ends; style ca. 4 mm, 2-fid; stigmas enlarged, ca. 3 mm. Fruit
oblong, 4–6 × 2–3 cm, smooth, rounded at both ends. Seeds
broadly ovate, 6–7 × ca. 5 mm, smooth, base obtuse-rounded.
Fl. and fr. Apr–Oct.
Forest margins on mountain slopes; 1200–1800 m. Yunnan [Bhutan, NE India, Laos, Myanmar, Thailand, Vietnam].

23

5. Thladiantha dubia Bunge, Enum. Pl. Chin. Bor. 29. 1833.
赤瓟 chi bao
Plants climbing, herbaceous, pale yellow pubescent-hirsute. Roots tuberous. Stems robust, angular-sulcate. Petiole robust, 2–6 cm; leaf blade broadly ovate-cordate, 5–8 × 4–9 cm,
both surfaces scabrous, hirsute on veins, basal lateral veinsalong margin of blade base, base cordate, margin irregularly
dentate, apex acute or shortly acuminate; sinus deep, suborbicular or semicircular, 1–1.5 cm deep, 1.5–3 cm wide. Tendrils
filiform, simple. Male flowers solitary or several congested on
short rachis into pseudoraceme; pedicel filiform, 1.5–3.5 cm,
pubescent; calyx tube very short, subrotate, 3–4 × 7–8 mm; segments reflexed, lanceolate, 12–13 × 2–3 mm; stamens 4 connected in pairs and 1 free, 2–2.5 mm, puberulent; anthers ovoid,ca. 2 mm; rudimentary ovary semiglobose. Female flowers solitary; pedicels slender, 1–2 cm, pubescent; calyx and corolla as
in male flowers; staminodes 5; ovary oblong, 5–8 mm, densely
pale yellow pubescent; style 3-fid; stigma enlarged, reniform,
2-lobed. Fruit orange-yellow, ovoid-oblong, 4–5 cm × ca. 28mm, pubescent, with 10 longitudinal striae, base slightly attenuate. Seeds black, ovate, 4–4.3 × 2.5–3 mm, ca. 1.5 mm thick,smooth. Fl. Jun–Aug, fr. Aug–Oct.
Forest margins, valleys; 300–1800 m. Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Ningxia, Shaanxi, Shandong, Shanxi, N Sichuan[Korea; introduced in Japan].
The fruit and roots are used medicinally.

6. Thladiantha sessilifolia Handel-Mazzetti, Symb. Sin. 7:
1061. 1936.
短柄赤瓟 duan bing chi bao
Vines, herbaceous. Roots tuberous. Stem and branches
slender, at first sparsely puberulent, glabrescent. Petiole very
short, 5–15(–40) mm, pubescent or subglabrous; leaf blade
lanceolate or ovate-lanceolate, 8–16 × 2–4.5 cm, papery, abaxially densely pubescent, adaxially scabrous, subglabrous, basecordate, margin sparsely dentate, apex acuminate; sinus closed,
basal lateral veins not along blade base. Tendrils filiform, usually simple. Male flowers 2–5 in pedunculate short raceme,
sometimes solitary; bract ovate, ca. 2 mm; pedicels slender, 5–
10 mm, pubescent or subglabrous; calyx tube campanulate, ca.
3 × 5 mm, sparsely pubescent; segments lanceolate, ca. 5 ×
1.5 mm, 3-veined, apex subulate-acuminate; corolla segments
ovate, ca. 15 × 7–8 mm, often 5-veined, apex acuminate; stamens inserted on limb of calyx tube; filaments ca. 3 mm; anthers elliptic, 2–3 mm; rudimentary ovary yellow, semiglobose.
Female flowers solitary; pedicel slender, 2–3 cm, puberulent or
subglabrous; calyx and corolla as in male flowers; ovary oblong, 15–22 × 5–10 mm, densely yellow-brown villous, apex
attenuate; style slender, 3-fid; stigma enlarged, reniform, 2lobed; staminodes 2–3 mm. Fruit ovoid-oblong. Fl. May–Aug,
fr. Aug–Nov.
● Thickets or forests on mountain slopes, wetlands on ditch sides.
SW Sichuan, Yunnan.
The tuberous roots are used medicinally.

1a. Petiole very short, 5–15 mm, pubescent................................................................ 6a. var. sessilifolia

CUCURBITACEAE

24沧源赤瓟 cang yuan chi bao

sulcate, pubescent and white setose at first, later glabrescent.
Petiole 4–8 cm; leaf blade broadly ovate-cordate, 6–12 × 4–10
cm, undivided, abaxially pubescent, adaxially scabrous, densely
setose, margin dentate, apex shortly acuminate or acute; basal
sinus open, semicircular, 5–15 mm deep, 10–25 mm wide.
Tendrils filiform, simple or 2-fid. Male flowers dimorphic, solitary one larger, smaller ones in a raceme; pedicel slender, 10–18mm, pubescent; calyx pubescent; tube broadly campanulate, 3–
4 mm; segments lanceolate, 4–6 mm, 3-veined, apex acuminate; corolla segments ovate, 10–12 × 5–6 mm, apex acute;filaments 2–2.5 mm; anthers elliptic, 3–3.5 mm. Female flowers solitary or binary; pedicels 5–10 mm; calyx and corolla as inmale flowers; corolla segments obovate, 20–25 × ca. 10 mm,
apex slightly obtuse; staminodes 5, sparsely puberulent, apex
slightly enlarged; ovary fusiform, densely villous; style 3-fid,
ca. 2 mm; stigma enlarged, 2-lobed.

Petiole longer, 20–40 mm, subglabrous. Tendrils simple or
2-fid.

xi.

1b. Petiole longer, 20–40 mm, subglabrous................................................................... 6b. var. longipes
6a. Thladiantha sessilifolia var. sessilifolia
短柄赤瓟(原变种) duan bing chi bao (yuan bian zhong)
Petiole very short, 5–15 mm, pubescent; leaf blade lanceolate or ovate-lanceolate. Tendrils simple.
● Thickets on mountain slopes, wetlands on ditch sides. SW Sichuan.
The tuberous roots are used medicinally.

6b. Thladiantha sessilifolia var. longipes A. M. Lu & Zhi Y.
Zhang, Bull. Bot. Res., Harbin 1(1–2): 77. 1981.

● Forests; ca. 1000 m. Yunnan.

7. Thladiantha villosula Cogniaux in Engler, Pflanzenr. 66(IV.
275. I): 44. 1916.
长毛赤瓟 chang mao chi bao
Thladiantha villosula var. nigrita A. M. Lu & Zhi Y.
Zhang.
Vines, climbing, herbaceous, densely glandular-villous andsparsely multicellular-setose. Stem much branched; branches
slender. Petiole slender, 3–6 cm; leaf blade adaxially dark green,
ovate-cordate, broadly ovate-cordate, or suborbicular, 6–12 ×
5–10 cm, membranous, abaxially pubescent, basal lateral veinsalong margin of blade base, base cordate, margin dentate, apex
shortly acuminate; sinus rounded, 1–2 cm deep, 1–1.5 cm wide.
Tendrils filiform, simple. Male flowers usually 2–7 in a raceme;
peduncle 1–3 cm, usually bearing leaflike bract; bract 1–1.5 ×
ca. 1 cm, densely pubescent, margin dentate, apex acuminate;pedicel 1–2.5 cm; calyx tube broadly campanulate; segments
yellow-green, narrowly lanceolate, 4–6 × ca. 1.5 mm, 3-veined;
corolla segments ovate or narrowly ovate, 12–15 × 6–8 mm, 5veined, apex obtuse; stamens 4 connected in pairs and 1 free;filaments filiform, 2–3 mm; anthers oblong, ca. 2 mm; rudimentary ovary semiglobose. Female flowers solitary; pedicels
slightly robust; calyx segments narrowly lanceolate, 5–6 × ca.
1 mm, 3-veined, apex acuminate; corolla segments narrowly
ovate, ca. 20 × 15 cm, 5-veined, apex acuminate; staminodes
glandular; ovary narrowly oblong, 15–18 × 3–4 mm, densely
pale yellow glandular villous; style stout, 3-fid; stigma enlarged, reniform, 2-lobed. Fruiting pedicel stout, 2–3 cm; fruit
red-brown, oblong, to 7 × ca. 3.5 cm, both surfaces reticulate,base obtuse, apex attenuate. Fl. and fr. summer.
● Ditch sides in forests or thickets; 2000–2800 m. Gansu, Guizhou, S Henan, W Hubei, S Shaanxi, Sichuan.

● Mountain slopes, roadsides; 1800–2400 m. E Shaanxi, S Shan-

9. Thladiantha nudiflora Hemsley, J. Linn. Soc., Bot. 23: 316.
1887.
南赤瓟 nan chi bao
Plants densely pubescent-hispid. Roots tuberous. Stems
herbaceous, angular-sulcate. Petiole stout, 3–10 cm; leaf blade
ovate-cordate, broadly ovate-cordate, or suborbicular-cordate,
5–15 × 4–12 cm, abaxially densely pale yellow pubescent,adaxially densely setose, basal lateral veins along blade base,margin serrate, apex acuminate or acute; basal sinus open or
sometimes closed, 2–2.5 cm deep, 1–2 cm wide. Tendrils
densely hispid, 2-fid. Male raceme many flowered; peduncle
filiform, 4–8 cm, densely pubescent; pedicels filiform, 1–1.5
cm; calyx densely villous; tube broadly campanulate, 5–6 mm
in diam.; segments ovate-lanceolate, 5–6 × ca. 2.5 mm, 3veined, apex acute; corolla segments ovate-oblong, 12–16 × 6–
7 mm, 5-veined, apex acute or obtuse; filaments puberulent, ca.
4 mm; anthers ovoid-oblong, ca. 2.5 mm. Female flowers solitary; pedicels slender, 1–2 cm, villous; calyx and corolla as inmale flowers; ovary narrowly oblong, 12–15 × 4–5 mm,
densely villous-hispid, base obtuse-rounded, apex attenuate;style 3-fid; stigma enlarged, reniform, 2-lobed; staminodes 5,
ca. 1.5 mm. Fruiting pedicel robust, 2.5–5.5 cm; fruit red or
red-brown when mature, oblong, 4–5 × 3–3.5 cm, glabrescent,apex obtuse. Seeds ovate or broadly so, ca. 5 × 3.5–4 mm, 1–
1.5 mm thick, reticulate, base rounded, apex acute. Fl. May–
Aug, fr. Aug–Nov.
Forest margins on mountain slopes. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi,
Shaanxi, Sichuan, Taiwan, Zhejiang [Philippines].

1a. Male flowers ebracteate .......................... 9a. var. nudiflora
1b. Male flowers with leaflike bracts ........... 9b. var. bracteata
9a. Thladiantha nudiflora var. nudiflora

8. Thladiantha dimorphantha Handel-Mazzetti, Oesterr. Bot.
Z. 83: 235. 1934.

南赤瓟(原变种) nan chi bao (yuan bian zhong)山西赤瓟 shan xi chi bao

Cucumis courtoisii H. Léveillé; Thladiantha formosana
Hayata; T. harmsii Cogniaux; T. nudiflora var. macrocarpa Z.
Zhang; T. nudiflora var. membranacea Z. Zhang.

Vines, climbing, herbaceous. Stem and branches angular-

CUCURBITACEAE

Male flowers ebracteate.
Forest margins on mountain slopes. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi,
Shaanxi, Sichuan, Taiwan, Zhejiang [Philippines].

9b. Thladiantha nudiflora var. bracteata A. M. Lu & Zhi Y.
Zhang, Bull. Bot. Res., Harbin 1(1–2): 84. 1981.
西固赤瓟 xi gu chi bao
Male flowers with leaflike bracts; bracts broadly ovatecordate, 10–12 × ca. 5 mm, pubescent, margin dentate.
● Gansu (Xigu).

10. Thladiantha longisepala C. Y. Wu, A. M. Lu & Zhi Y.
Zhang, Bull. Bot. Res., Harbin 1(1–2): 86. 1981.
长萼赤瓟 chang e chi bao
Herbs, climbing. Stem and branches angular-sulcate, atfirst puberulent and sparsely setose, glabrescent. Petiole slender,
4–7 cm, puberulent; leaf blade ovate-cordate or broadly ovatecordate, 8–14 × 6–10 cm, membranous, abaxially puberulent,
adaxially densely setose, base cordate, margin serrate, apex
shortly acuminate; sinus 1–2 cm deep, 1.5–3 cm wide. Tendrils
robust, elongate, 2-fid. Male panicle much branched, 5–13 cm,
sparsely pubescent, usually with a leaflike bract at base ofbranch; bract ovate, 15–25 × 10–15 mm, apex acuminate orlong acuminate. Male flowers: pedicel slender, 1–4 cm, pubescent; calyx tube campanulate; segments broadly lanceolate,
12–15 × 3–4 mm, distinctly 3-veined, apex acuminate, reflexed;
corolla segments narrowly ovate, 20–22 × 8–9 mm, 5-veined,
apex acuminate; filaments ca. 4 mm, puberulent; anthers ovoid,ca. 3 mm; rudimentary ovary semiglobose. Female flowers: 2–5
on a short peduncle; pedicels 3–6 cm; ovary densely setose andpubescent, oblong, ca. 1 cm. Fruiting peduncle robust, 3–6 cm,
puberulent; fruit oblong, scabrous, yellow-brown pubescent andsetose, apex and base obtuse. Fl. and fr. Jun–Oct.
● Thickets on mountain slopes and riversides; 2400–3500 m. NW
Yunnan.

11. Thladiantha setispina A. M. Lu & Zhi Y. Zhang, Bull. Bot.
Res., Harbin 1(1–2): 87. 1981.
刚毛赤瓟 gang mao chi bao
Herbs, climbing. Stems and branches sparsely puberulent,ultimately glabrous, smooth, angular-sulcate. Petiole slender,
3.5–5.5 cm, glabrous; leaf blade ovate-cordate, 6–8(–14) × 4–
6(–10) cm, membranous or papery, abaxially densely pale yellow pubescent, adaxially setose, basal lateral veins along blade
base, base cordate, margin serrulate, apex acuminate; sinus
open, semicircular, 1–1.5 cm deep, 1–2 cm wide. Tendrils filiform, glabrous, 2-fid. Male flowers numerous in a ca. 17 cm
panicle; peduncle robust, sparsely pubescent, usually with abract at base of branches; bract 2–2.5 × 1.5–2 cm; pedicel 1–1.5
cm; calyx tube campanulate; segments triangular-lanceolate, 5–
6 × ca. 2.5 mm, 3-veined, apex acuminate; corolla segments
oblong, 18–20 × 6–8 mm, glabrous, 5-veined, apex acuminate;filaments ca. 4 mm, puberulent; anthers elliptic, ca. 3 mm. Female flowers solitary, or 3–5 on a 3–4 cm peduncle; pedicel
slender, 3–5 cm, sparsely puberulent; calyx tube campanulate;segments ca. 7.5 mm, ca. 2.5 mm wide at base, glabrous, 3-25veined, apex acuminate; corolla segments 4–4.5 × 1.8–2 cm;
ovary oblong, 10–12 × 6–7 mm, densely yellow-brown setose;style 3-fid; stigma enlarged, reniform, 2–2.5 mm, 2-lobed.
Fruiting peduncle robust, 3–6 cm, glabrescent; fruit blackbrown, oblong, 3–3.5 × 2–2.5 cm, scabrous, yellow-brownsetose, apex and base obtuse. Seeds yellow-brown when dry,narrowly ovate, 6–7 × 3–3.5 mm, ca. 1.5 mm thick, smooth,base subrounded, apex obtuse. Fl. and fr. Jun–Oct.
● Mountain slopes, roadsides; ca. 3000 m. W Sichuan, E Xizang.

12. Thladiantha lijiangensis A. M. Lu & Zhi Y. Zhang, Bull.
Bot. Res., Harbin 1(1–2): 88. 1981.
丽江赤瓟 li jiang chi bao
Thladiantha cinerascens C. Y. Wu ex A. M. Lu & Zhi Y.
Zhang; T. henryi Hemsley var. subtomentosa Handel-Mazzetti;
T. lijiangensis var. latisepala A. M. Lu & Zhi Y. Zhang.
Herbs, climbing. Root robust. Stem and branches distinctly angular-sulcate, at first pubescent, glabrescent. Petiole
slender, 2–6 cm, pubescent and sparsely setose; leaf blade ovatecordate, 4–11 × 3–7 cm, membranous, abaxially glaucous andsetose, adaxially pale yellow pubescent, base cordate, margindentate, apex acuminate; sinus 1–2 cm deep, 1–2.5 cm wide.
Male panicle much branched, with many flowers; peduncle 7–
16 cm, usually with a leaflike bract at base of branches; bract
ovate, 1–2 cm; pedicels filiform, 1–2 cm, puberulent; calyx
pubescent; tube campanulate; segments triangular-lanceolate,ca. 5 × 1.5–2 mm, 1-veined, apex shortly acuminate; corollasegments ovate, 12–15 × ca. 6 mm, 5-veined, apex acute; filaments ca. 2.5 mm; anthers narrowly ovoid, ca. 2.5 mm; rudimentary ovary semiglobose, ca. 1.5 mm in diam. Female flowers larger than male flowers, solitary or binary, sometimes 3 atapex of 15–25 mm peduncle; pedicels 4–6 cm; calyx segments
lanceolate, ca. 6 × 1.5 mm, sparsely puberulent, 1-veined; corolla segments ovate, 30–35 × ca. 13 mm, puberulent, 5-veined,
apex acuminate; ovary ellipsoid, 10–13 × 6–7 mm, densely
yellow hispidulous, base subrounded, apex truncate; style ca. 4
mm, 3-fid; stigma reniform, 2–2.5 mm wide, 2-lobed. Fruiting
peduncle 3–4 cm, puberulent; fruit 3–4 × 2–2.5 cm. Seeds grayblack, broadly ovate, ca. 5.5 × 4 mm, smooth. Fl. and fr. May–
Oct.
● Forests, mountain slopes; 2400–3200 m. NW Yunnan.

13. Thladiantha oliveri Cogniaux ex Mottet, Rev. Hort. 75:
473. 1903.
鄂赤瓟 e chi bao
Thladiantha glabra Cogniaux.
Herbs, climbing. Stem and branches slender, glabrous, angular-sulcate. Petiole 5–15 cm, subglabrous; leaf blade broadly
ovate-cordate, 10–20 × 8–18 cm, membranous, abaxiallysparsely setose, adaxially scabrous, basal lateral veins alongbase of sinus, margin dentate, apex acute or acuminate; sinus
open, 1.5–3 cm deep. Tendrils glabrous, 2-fid. Male flowers
numerous at apex of peduncle; peduncle over 20 cm; pedicels
5–10 mm, slender; calyx tube broadly campanulate; segments
reflexed, linear, 7–9 mm, 1-veined, apex acuminate; corollasegments ovate-oblong, 18–22 × 7–9 mm, 5-veined, apex acuminate; filaments 2–3 mm, sparsely puberulent; anthers ovoid-

CUCURBITACEAE

26

oblong, ca. 2 mm; rudimentary ovary globose. Female flowers
solitary or 2–4 on a 10–15 mm peduncle; pedicels 2–4 cm,
subglabrous; calyx segments reflexed, linear, 10–12 mm; corolla segments 2–4 × ca. 1.2 cm; staminodes linear, ca. 2 mm;ovary ovoid, 10–12 mm, base truncate, apex attenuate. Fruitovoid, 3–4 × 2–2.5 cm, glabrous, smooth, base truncate, apex
attenuate. Seeds ovate, compressed, 5–6 × 3–3.5 mm, ca. 1.5mm thick, base obtuse, apex attenuate. Fl and fr. May–Oct.
● Thickets, roadsides, or wetlands in river valleys; 600–2100 m.
Gansu, Guizhou, NW Hubei, S Shaanxi, E and S Sichuan.

14. Thladiantha montana Cogniaux in Engler, Pflanzenr.
66(IV. 275. I): 48. 1916.
山地赤瓟 shan di chi bao
Herbs, climbing, glabrous. Stem and branches angular-sulcate. Petiole slender, 8–10 cm; leaf blade ovate-cordate or
broadly ovate-cordate, 10–16 × 8–12 cm, submembranous or
thickly papery, adaxially scabrous, base cordate, margin dentate, apex acuminate; sinus suborbicular. Tendrils 2-fid. Male
flowers 5–10 in raceme or panicle, sometimes 3 or 4 inflorescences in same leaf axil; rachis slender, 6–10 cm; pedicels filiform, 5–10 mm; calyx tube cupular; segments triangular-lanceolate, ca. 4 × 1 mm, 1-veined, apex obtuse; corolla segments
narrowly ovate, 8–12 × ca. 5 mm, 5-veined, apex acuminate;stamens inserted on tube; filaments 4 connected in pairs and 1
free, 2–3 mm, sparsely pubescent; anthers oblong, ca. 2 mm;
rudimentary ovary subglobose, ca. 2 mm in diam. Female flowers solitary, or 2 or 3 on a 1–2.5 cm peduncle; pedicels slender,
1–4 cm, glabrous; calyx and corolla as in male flowers; staminodes 2.5–3 mm, glandular, with 3 membranous scales; scales
oblong, 3–4 mm, apex obtuse; ovary narrowly oblong, 10–15 ×
3–4 mm, smooth, subglabrous, base truncate, apex attenuate;style slender, 5–6 mm, 3-fid; stigma suborbicular, 4.5–5 mmwide, 2-lobed. Fruiting peduncle smooth; fruit oblong, 4–6 × 3–
4 cm, both ends obtuse-rounded. Seeds narrowly ovate, 4–5 ×
2–3 mm, ca. 1.5 mm thick, base rounded. Fl. Aug–Oct, fr. Oct–
Nov.
● Forests, mountain slopes, valley sides; 800–3200 m. Yunnan.

15. Thladiantha maculata Cogniaux in Engler, Pflanzenr.
66(IV. 275. I): 49. 1916.
斑赤瓟 ban chi bao
Herbs, climbing. Roots tuberous. Stem and branches slender, angular, puberulent or subglabrous. Petiole slender, 4–9cm, puberulent; leaf blade broadly ovate-cordate, 7–13 × 5–10
cm, membranous, abaxially puberulent, adaxially setose, basal
lateral veins along base of blade, base cordate, margin dentate
or irregularly serrate, apex shortly acuminate; sinus open, semicircular. Tendrils filiform, simple, subglabrous or sparsely puberulent. Male flowers in a raceme of 3–6(–8) flowers; rachis
slender, 3–4 cm, puberulent; pedicels filiform, 12–25 mm,smooth; calyx tube broadly campanulate; segments narrowly
triangular-lanceolate, ca. 4 mm, pubescent, 3-veined, apex acuminate; corolla segments ovate, 12–15 × ca. 7 mm, 5-veined,
apex acute or shortly acuminate; filaments ca. 3 mm, puberulent; anthers oblong, ca. 2 mm; rudimentary ovary semiglobose.
Female flowers solitary; pedicels filiform, 25–35 mm, pubescent; calyx segments linear-subulate, 7–8 mm, puberulent; co-rolla segments as in male flowers; ovary oblong or narrowly
fusiform, ca. 10 mm, densely pale yellow pubescent, base truncate, apex acuminate; style 3-fid; stigma enlarged, reniform, 2lobed. Fruiting peduncle 3–4 cm, puberulent, glabrescent; fruitorange-red, fusiform, 5–7 × 2–2.8 cm, smooth, subglabrous,
base attenuate, apex acuminate, rostrate. Seeds narrowly ovate,
4–5 × ca. 2.5 mm, ca. 1.5 mm thick, smooth. Fl. May–Aug, fr.
Oct.
● Valleys, forests; 500–1800 m. Henan, W Hubei.

16. Thladiantha punctata Hayata, J. Coll. Sci. Imp. Univ.
Tokyo 30(1): 119. 1911.
台湾赤瓟 tai wan chi bao
Herbs, climbing; whole plant subglabrous. Stems andbranches with obvious longitudinal striae. Petiole 3–12 cm,
sometimes sparsely shortly setose in upper part; leaf blade
abaxially pale green, adaxially dark green, narrowly ovate or
narrowly ovate-lanceolate, 8–16(–20) × 6–10(–12) cm, membranous, glabrous, lateral veins at base not bordering sinus,
margin dentate or sinuate-dentate, apex acuminate; basal sinus
semicircular, 1–2 cm deep, 0.5–1.5 cm wide. Tendrils simple.
Male flowers: inflorescences racemose or paniculate, rarely
flowers solitary; peduncles slender, 4–8 cm; pedicels filiform,
0.5–1 cm; calyx tube broadly campanulate, 5–6 mm; segments
lanceolate, ca. 8 mm, ca. 2 mm wide at base, 3–5-veined,
acuminate at apex; stamens 4 in pairs and 1 free; filaments ca. 4
mm, base pubescent; anthers oblong, 2–3 mm; rudimentary
ovary semiglobose, 2–3 mm in diam., with 3 membranous
scales at base; scales yellow, ca. 3 mm. Female flowers usuallysolitary, rarely 2 at apex of peduncle; peduncles ca. 1 cm;
pedicels 2–5 cm, glabrous; calyx and corolla as in male flowers
but corolla usually larger than in male flowers; segments 2–2.5
× 1.1–1.2 cm, acuminate at apex; staminodes 4 in pairs and 1
free, ca. 1 mm; ovary ovoid, 1–1.3 × 0.4–0.6 cm, slightly pubescent, glabrescent, base rounded, apex attenuate; styles 2–3
mm, 3-fid; stigma reniform, expanded, 1.5–2 mm in diam.
Fruiting pedicel 3–7 cm, glabrous; fruit ovoid or oblong, 3–5 ×
2–3.5 cm, smooth, base obtuse-rounded, apex mucronate. Seeds
brown, broadly ovoid, 5–6 × 3–4 mm, ca. 1 mm thick, baserounded, apex slightly narrowed. Fl. May–Jun, fr. Jul–Aug.
● Mountain slopes, forests, wet places; 600–900 m. Anhui, Fujian, Jiangxi, Taiwan, Zhejiang.

17. Thladiantha dentata Cogniaux in Engler, Pflanzenr. 66(IV.
275. I): 44. 1916.
齿叶赤瓟 chi ye chi bao
Herbs, climbing, stout; whole plant almost glabrous. Stemand branches smooth, angular-sulcate. Petiole robust, 5–16 cm;
leaf blade ovate-cordate or broadly ovate-cordate, 12–20 × 8–
12 cm, abaxially glabrous, adaxially scabrous, basal lateralveins not along margin of blade, margin dentate, apex shortly
acuminate; basal sinus open or sometimes closed. Tendrils 2fid. Male flowers: inflorescence racemose or paniculate; rachis
slender, 8–12 cm; pedicels filiform, 1–1.5 cm; calyx tube cupular; segments oblong-lanceolate, ca. 5 × 1.5 mm, 3-veined,
apex obtuse; corolla segments ovate-oblong, ca. 12 × 5–6 mm,
3–5-veined, apex acute; stamens 4 in pairs and 1 free; filaments
ca. 4 mm; anthers elliptic, ca. 2 mm; rudimentary ovary semi-

CUCURBITACEAE

globose, ca. 2 mm in diam., with 3 scales at base; scales yellow,
ca. 3 mm. Female flowers solitary, or 2–5 at apex of 1–1.5 cm
peduncle; pedicels 3–6 cm, glabrous; calyx segments lanceolate, 4–5 × ca. 1.5 mm, 3-veined, apex acute; corolla segments
ovate-oblong, ca. 15 × 7–8 mm, 5-veined, apex acute; staminodes 5, 4 in pairs, other free, ca. 2.5 mm; ovary narrowly oblong, 13–16 × 4–6 mm, smooth, base slightly rounded or truncate, apex attenuate; style 2–3 mm, 3-fid; stigma reniform, 2lobed. Fruiting peduncle 2–3.5 cm; fruit narrowly ellipsoid or
narrowly ovoid, 3.5–6 × 2.5–3.5 cm, smooth, rounded at bothends, apex mucronate. Seeds yellow-white, narrowly ovate, ca.
6 × 3.5 mm, smooth, base rounded, apex slightly attenuate. Fl.
summer, fr. Oct–Nov.
● Roadsides, valley sides, thickets, mountain slopes; 500–2000 m.
Guizhou, W Hubei, Hunan, Sichuan.

18. Thladiantha capitata Cogniaux in Engler, Pflanzenr.
66(IV. 275. I): 51. 1916.
头花赤瓟 tou hua chi bao
Vines, climbing, herbaceous. Stem and branches angularsulcate, subglabrous. Petiole slender, 7–9 cm, glabrous; leaf
blade abaxially pale green, broadly ovate or broadly ovate-triangular, 10–15(–20) × 8–12(–15) cm, membranous, both surfaces glabrous, adaxially scabrous, basal lateral veins along
margin of sinus, margin sparsely dentate, apex acuminate; basal
sinus 1.5–3 cm deep, 2–3.5 cm wide. Tendrils glabrous, 2-fid.
Male flowers at apex of peduncle in congested subumbel, umbelliform raceme, or pseudocapitulum, usually 8–15-flowered;peduncle (2–)5–11(–17) cm, glabrous, sometimes with a bract;
bract ovate, 1.5–3 × 1–2.8 cm; pedicels filiform, 4–8 mm; calyx
tube obconical, ca. 5 mm; segments linear-lanceolate, ca. 10 ×
1.5 mm, abaxially puberulent, adaxially glabrous, 1-veined,
apex acute; corolla segments narrowly ovate, 10–12 × 5–6 mm,
adaxially pubescent, 5-veined, apex acute; filaments 4–4.5 mm,
puberulent; anthers ovoid-oblong, 2–2.2 mm. Female flowers
solitary, or 2 or 3 flowers at apex of peduncle; pedicels 3–5 cm
when flowers solitary; calyx segments linear-lanceolate, 8–12 ×
1.5–1.8 mm, 1-veined; corolla segments oblong, 15–28 × 1–12mm, 5-veined, apex acuminate; ovary narrowly oblong, 15–18mm, smooth, base subtruncate, apex attenuate; style slender, ca.
5 mm, 3-fid; stigma enlarged, reniform, ca. 2 mm wide. Fruit
oblong, ca. 4 × 2.5 cm. Fl. and fr. May–Nov.
● Forest margins, thickets, mountain slopes; 1000–2700 m. W Sichuan.

19. Thladiantha davidii Franchet, Nouv. Arch. Mus. Hist.
Nat., sér. 2, 8: 243. 1885 [“davidi”].
川赤瓟 chuan chi bao
Thladiantha legendrei Gagnepain.
Herbs, climbing. Stems and branches glabrous, longitudinally angular-sulcate. Petiole 6–8 cm, glabrous; leaf blade
ovate-cordate, 10–20 × 6–12 cm, membranous, abaxially glabrous, adaxially densely setose, basal lateral veins along marginof sinus, margin dentate, apex acuminate; basal sinus rounded,sometimes closed. Tendrils 2-fid. Male flowers 10–20 at apexof peduncle in congested subumbelliform raceme; peduncle 10–
20 cm, sometimes with a leaflike bract; bract 15–20 × 10–1327mm; pedicels slender, 3–6 mm or sometimes to 15 mm; calyx
tube obconical; segments lanceolate-oblong, 10–12 × ca. 2.5mm, abaxially puberulent, distinctly 3-veined, apex obtuse or
acute; corolla segments ovate, ca. 15 × 9 mm, adaxially glandular pubescent, 5-veined, apex obtuse, base of corolla with 2
yellow scales; filaments 3–4 mm, sparsely puberulent; anthers
elliptic, ca. 1.5 mm. Female flowers solitary, or 2 or 3 at apex ofpeduncle; peduncle ca. 1(–3) cm; pedicels 1.5–3 cm; calyx tube
obconical; segments lanceolate-oblong, 10–15 × ca. 3 mm,
distinctly 3-veined, base slightly narrowed, apex obtuse; corollasegments oblong, 2.5–2.7 × 1–1.2 cm, 5-veined, apex acuminate; ovary narrowly oblong, ca. 15 × 5–6 mm, smooth, subglabrous, base truncate, apex slightly attenuate; style 3-fid; stigma 2-lobed, reniform. Fruit oblong, 3–4.5 × 2–2.4 cm, base andapex obtuse. Seeds yellow-white, ovate, compressed, 3–4 × ca.
2.5 mm, smooth. Fl. and fr. May–Nov.
● Roadsides, valleys, thickets; 1100–2100 m. Guizhou, W Sichuan.

20. Thladiantha palmatipartita A. M. Lu & C. Jeffrey, Novon
10: 398. 2000.
掌叶赤瓟 zhang ye chi bao
Herbs, scandent. Stems sulcate-angular, 1–3 m, puberulent
or glabrescent. Tendrils simple, pubescent. Petiole 3–7 cm, puberulent; leaf blade broadly ovate or orbicular in outline, 8–17
× 9–16 cm, papery, abaxially densely pubescent, adaxiallydensely punctate, palmately deeply 5-lobed; lobes oblong, margin irregularly dentate, apex acute or acuminate. Flowers unisexual. Male flowers: panicle short, many flowered; peduncle
2.5–3 cm, robust, puberulent; pedicels slender, filiform, 5–12mm; calyx tube cupular, pubescent; segments 5, broadly lanceolate, 1.2–1.5 × ca. 0.5 cm, 3-veined, apex acuminate; corollacampanulate; segments 5, narrowly oblong or lanceolate-oblong, 1.5–1.8 × ca. 0.6 cm, 3-veined, apex acuminate; filaments
ca. 2 mm; anthers ca. 3 mm, 1-celled. Female flowers solitaryor paired; calyx and corolla as in male flowers; ovary narrowly
oblong, 1.5–2 cm, villous; style columnar, apically 3-fid; stigmas dilated, 2-lobed. Fruit oblong, ca. 7.5 × 3.5 cm, smooth,base attenuate, apex rounded. Seeds numerous, horizontal, ca. 6
× 3 mm, smooth. Fl. Jun, fr. Sep.
● Open situations in scrub; ca. 3000 m. Yunnan.

21. Thladiantha pustulata (H. Léveillé) C. Jeffrey ex A. M.
Lu & Zhi Y. Zhang, Bull. Bot. Res., Harbin 1(1–2): 74. 1981.
云南赤瓟 yun nan chi bao
Herbs, climbing. Roots tuberous. Stem and branches angular-sulcate, pubescent and villous. Petiole slender, 3–7 cm, pubescent and villous; leaf blade ovate-cordate, 6–11 × 4–9 cm,
abaxially puberulent and setose on veins, adaxially densely
setose, scabrous, basal lateral veins along margin of sinus, basecordate, margin irregularly serrate, apex shortly acuminate;sinus open, semicircular, 15–20 mm deep, 10–20 mm wide.
Tendrils simple, pubescent. Male flowers 3–6 at apex of peduncle in sparse racemes; peduncle 2–5 cm, pubescent; pedicels
filiform, 2–4 cm; calyx tube campanulate, ca. 4 × 4–5 mm;segments slightly reflexed, linear or linear-lanceolate, 4–12mm, 3-veined, apex acuminate; corolla segments ovate-lanceolate, ca. 15 × 6 mm, apex acute; filaments filiform, 3–4 mm,

CUCURBITACEAE

28

puberulent; anthers oblong, ca. 2 mm. Female flowers solitary,larger than male flowers; pedicel slender, pubescent; calyx segments linear, 8–9 mm; corolla segments ovate-lanceolate, ca. 20
× 7 mm, distinctly 3–5-veined, apex shortly acuminate; ovary
ovoid, ca. 7 × 6 mm, pubescent and villous, apex shortly rostrate; style ca. 5 mm, 3-fid; stigma enlarged, 2-lobed, reniform.
Fruiting pedicel 2–3 cm, sparsely pubescent; fruit ovoid, 2–3
cm in diam., verrucose. Seeds broadly ovate, 6–7 × ca. 3.5 mm,
ca. 1.5 mm thick, punctate, base slightly obtuse. Fl. and fr. Apr–
Nov.
● Valleys, thickets, forest margins; 1100–2600 m. Chongqing,
Guizhou, Sichuan, Yunnan.

1a. Leaf blade lobed; calyx segments
4–8 mm ................................................. 21a. var. pustulata
1b. Leaf blade dentate, undivided;
calyx segments 9–12 mm ........... 21b. var. jingfushanensis
21a. Thladiantha pustulata var. pustulata

base of blade, base cordate, margin dentate, sometimes irregularly triangular-serrate, apex acute or shortly acuminate; sinus
open, semicircular, 1–3 cm deep, 1–2 cm wide. Tendrils filiform, 2-fid or simple. Male flowers 6–10 at apex of peduncle inraceme or panicle; peduncle 5–12 cm, puberulent; pedicel 1–3
cm, puberulent; calyx tube campanulate, 5–7 mm in diam.; segments lanceolate, 10–12 mm, puberulent, 1-veined, apex acuminate; corolla segments oblong-elliptic or oblong, ca. 20 × 6
mm, 5-veined, apex shortly acuminate or acute; stamens 4 connected at base in pairs and 1 free; filaments 4–5 mm, puberulent, with 3 scales at base; anthers oblong, ca. 4 mm; rudimentary ovary absent. Female flowers solitary, or 2 or 3 to many ina raceme; pedicels 2–6 cm; calyx and corolla as in male flowers; staminodes 5, ca. 2.5 mm; ovary narrowly ovoid or ovoidoblong, verrucose-rugose; style 3–4 mm; stigma enlarged, reniform, 2-lobed. Fruiting pedicel 6–9 cm; fruit ellipsoid, 5–10 ×
3–4 cm, rugose, base attenuate at pedicel to 10 mm. Seeds
narrowly ovate, 5–6 × 2–2.5 mm, smooth. Fl. and fr. Jun–Nov.

云南赤瓟(原变种) yun nan chi bao (yuan bian zhong)● Thickets, forests, roadsides; 1100–2000 m. W Hubei, W Hunan,
S Shaanxi, E Sichuan.

Melothria pustulata H. Léveillé, Cat. Pl. Yun-Nan, 65.
1916; M. mairei H. Léveillé; Thladiantha yunnanensis Gagnepain.

23. Thladiantha longifolia Cogniaux ex Oliver, Hooker’s Icon.
Pl. 23: t. 2222. 1892.
长叶赤瓟 chang ye chi bao

Leaf blade lobed. Calyx segments 4–8 mm.
● Valleys, thickets; 1500–2600 m. Guizhou, Yunnan.

21b. Thladiantha pustulata var. jingfushanensis A. M. Lu &J. Q. Li, Acta Bot. Yunnan. 14: 134. 1992.
金佛山赤瓟 jin fo shan chi bao
Leaf blade margin dentate, undivided. Calyx segments 9–
12 mm.
● Wetlands in valleys, forest margins; 1100–1800 m. Chongqing(Jinfo Shan), Sichuan.

22. Thladiantha henryi Hemsley, J. Linn. Soc., Bot. 23: 316.
1887.
皱果赤瓟 zhou guo chi bao
Thladiantha dictyocarpa Handel-Mazzetti; T. henryi var.
verrucosa (Cogniaux) A. M. Lu & Zhi Y. Zhang; T. verrucosa
Cogniaux.
Vines, climbing. Roots tuberous, enlarged. Stem andbranches longitudinally angular-sulcate, sparsely puberulent,glabrescent. Petiole slender, 4–12 cm, puberulent; leaf blade
broadly ovate-cordate, 8–16 × 7–14 cm, membranous, abaxiallypuberulent, adaxially white setose, basal lateral veins along

Vines, climbing. Stems and branches slender, angular, glabrous or sparsely pubescent. Petiole 2–7 cm, glabrous or shortly
pubescent; leaf blade ovate-lanceolate or narrowly ovate-triangular, 8–18 × 4–8 cm, membranous, abaxially subglabrous,
adaxially white setose, basal lateral veins not along base of
blade, base cordate, margin dentate, apex acute or shortly acuminate; sinus open, semicircular, 1.5–2 cm deep, 1.5–2.5 cm
wide. Tendril filiform, simple, glabrous. Male flowers 3–9(–12)
at apex of peduncle in a raceme; peduncle slender, 2–2.5 cm;
pedicels 1–2 cm, sparsely pubescent, glabrescent; calyx tube
shortly campanulate, ca. 0.6 cm wide at apex, pubescent onveins; segments triangular, 7–8 mm, 1-veined; corolla segments
oblong or elliptic, 1.5–2 × 0.8–1 cm, 5-veined, apex slightly obtuse; stamens 4 connected in pairs at base and 1 free; filaments
ca. 3 mm; anthers oblong, 2.5–3 mm. Female flowers solitary,or 2 or 3 flowers on short peduncle; pedicels 2–4 cm; calyx andcorolla as in male flowers; staminodes 5, subulate, ca. 1.5 mm,
4 in pairs, 1 free; ovary ovoid, acuminate at both apex and base,
base emarginate and with lobules, rugose; style 3-fid; stigma
enlarged, reniform. Fruit broadly ovoid, ca. 4 cm, verrucose,base slightly emarginate. Seeds ovate, 6–8 × 3–4.5 mm, 1–1.5
mm thick, apex obtuse. Fl. Apr–Jul, fr. Aug–Oct.
● Mountain slopes, forests, valleys, thickets; 1000–2200 m.
Guangxi, Guizhou, Hubei, Hunan, Sichuan.

12. MOMORDICA Linnaeus, Sp. Pl. 2: 1009. 1753.
苦瓜属 ku gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, annual or perennial, scandent or creeping. Tendrils unbranched or 2-fid. Petiole usually glandular; leaf blade suborbicular or ovate-cordate, palmately 3–7-lobed, rarely undivided, entire or dentate. Plants dioecious or monoecious. Male flowers solitary
or in a raceme; pedicel often with a large bract; bract orbicular-reniform; calyx tube short, campanulate or cupular; segments ovate,lanceolate, or oblong-lanceolate; corolla yellow or white, rotate or broadly campanulate, 5-lobed; segments obovate, oblong, or

CUCURBITACEAE

29

ovate-oblong; stamens (2 or)3(or 5); filaments short, free; anthers coherent at first, one 1-celled, other 2-celled; anther cells reflexed,straight, or curved; connective not produced; rudimentary ovary glandular or absent. Female flowers: ovary oblong or fusiform; styleelongate; stigmas 3, undivided or 2-lobed; ovules numerous, horizontal. Fruit ovoid, oblong, elliptic, or fusiform, undivided or 3valved, usually verrucose or spinescent. Seeds few or numerous, ovate or oblong, smooth or reticulate.
Forty-five species: most in tropical Africa, some cultivated in tropical regions; three species (one introduced) in China.

1a. Plants monoecious; bract on middle of male pedicel; leaf blade 5–7-partite; stamens 3; fruit fusiform or
cylindric, verrucose ..................................................................................................................................................... 1. M. charantia
1b. Plants dioecious; bract at apex of male pedicel.
2a. Petiole eglandular; anther cells conduplicate; fruit smaller than 5 cm, ovoid or ovoid-oblong, verrucose
or longitudinally sulcate ................................................................................................................................... 2. M. subangulata
2b. Petiole with 2–5 median glands; anther cells reflexed; fruit larger than 5 cm, ovoid, densely spinescent...................................................................................................................................................................... 3. M. cochinchinensis
1. Momordica charantia Linnaeus, Sp. Pl. 2: 1009. 1753.
苦瓜 ku gua
Cucumis argyi H. Léveillé; Momordica chinensis Sprengel; M. indica Linnaeus; M. sinensis Sprengel; Sicyos fauriei H.
Léveillé.
Plants annual, scandent, many branched; stem andbranches pubescent. Tendrils to 20 cm, puberulent, simple. Petiole slender, 4–6 cm, white pubescent at first, glabrescent; leaf
blade ovate-reniform or suborbicular, 4–12 × 4–12 cm, membranous, puberulent on veins, 5–7-partite; lobes ovate-oblong,veins palmate, margin crenate or irregularly lobed, apex obtuse
or acute; sinus semicircular. Plants monoecious. Male flowers
solitary in axils of leaves; pedicel slender, 3–7 cm, puberulent,with a median bract; bract reniform or orbicular, 5–15 mm,
entire, both surfaces puberulent; calyx segments ovate-lanceolate, 4–6 × 2–3 mm, white pubescent, apex acute; corolla yellow; segments obovate, 15–20 × 8–12 mm, pubescent, obtuse
or retuse; stamens 3, free; anther cells conduplicate. Female
flowers solitary; pedicel 10–12 cm, with a bract at base; ovary
fusiform, densely verrucose; stigmas expanded, 2-lobed. Fruitorange when mature, fusiform or cylindric, 10–20 cm, verrucose, 3-valved from apex. Seeds numerous, oblong, 15–20 ×
10–15 cm. Fl. and fr. May–Oct.
Commonly cultivated in China [pantropical, also cultivated intemperate and tropical regions].
The fruit is used as a vegetable; all parts of the plant are used
medicinally.

2. Momordica subangulata Blume, Bijdr. 928. 1826.
凹萼木鳖 ao e mu bie
Plants scandent. Stem and branches sulcate-angular, glabrous or puberulent at nodes. Petiole slender, 3–8 cm, glabrous,
eglandular; leaf blade ovate-cordate or broadly ovate-cordate,
6–13 × 4–11 cm, membranous, both surfaces puberulent at first,
glabrescent, base cordate, sinus subquadrate, margin dentate,usually undivided or sometimes 3–5-lobed, apex acute or acuminate. Tendrils filiform, simple. Plants dioecious. Male flowers solitary in leaf axil; pedicel slender, 5–8(–14) cm, puberulent, apex bracteate; bract orbicular-reniform, 10–30 × 10–40
mm, both surfaces puberulent, glabrescent, base retuse, apex
rounded to subacute; calyx segments ovate-oblong or lanceo-

late, 7–9 × 4–5 mm, pubescent, apex obtuse, retuse, broadly
rounded, or acute; corolla yellow; segments oblong or obovate,
20–30 × 15–25 mm, 5-veined, apex obtuse-rounded; stamens 3
or 5; filaments filiform, ca. 3 mm, glabrous; anthers 2.5–3 mm;anther cells conduplicate. Female flowers solitary; pedicel filiform, 5–10 cm, usually with a minute bract at base; calyx andcorolla as in male flowers; ovary ovoid-oblong or fusiform, 8–
12 × 2–4 mm, ridged or verrucose, base attenuate. Fruiting
pedicel slender, glabrous; fruit ovoid or ovoid-oblong, 3–7 ×
2.5–4 cm, longitudinally sulcate or with rows of tubercles, bothends attenuate. Seeds gray or yellow-brown, oblong, ovoid, or
globose, 7–14 × 5–8 mm, both surfaces slightly sculptured. Fl.
Jun–Aug, fr. Aug–Nov.
Mountain slopes, thickets, roadsides; 800–2500 m. Guangdong,
Guangxi, Guizhou, Yunnan [Bangladesh, India, Indonesia, Laos, Malaysia, Myanmar, Thailand, Vietnam].

1a. Plant slender; bract of male flower
10–20 × 10–20 mm; fruit 3–5 cm
with irregularly undulate longitudinalribs, glabrous or almost so ............ 2a. subsp. subangulata
1b. Plant more robust; bract of male
flower 20–30 × 20–40 mm; fruit
5–7 cm, with longitudinal rows offlattened tubercles or undulate ridges,
hispid-verrucose ................................... 2b. subsp. renigera
2a. Momordica subangulata subsp. subangulata
凹萼木鳖(原亚种) ao e mu bie (yuan ya zhong)Momordica eberhardtii Gagnepain; M. laotica Gagnepain.
Stems slender. Leaf blade puberulent at first on both surfaces, glabrescent. Bract of male flower 10–20 × 10–20 mm.
Calyx segments ovate-oblong, pubescent, apex obtuse, retuse.
Fruit ovoid or ovoid-oblong, 3–5 cm, longitudinally sulcate, notverrucose, glabrous or almost so. Fl. Jun–Aug, fr. Aug–Oct.
Mountain slopes, roadsides; 800–1500 m. Guangdong, Guangxi,
Guizhou, Yunnan [Indonesia, Laos, Malaysia, Myanmar, Thailand, Vietnam].

2b. Momordica subangulata subsp. renigera (Wallich ex G.
Don) W. J. de Wilde, Bot. Zhurn. 87: 147. 2002.
云南木鳖 yun nan mu bie

CUCURBITACEAE

30Momordica renigera Wallich ex G. Don, Gen. Hist. 3: 36.
1834.
Stem robust. Leaf blade abaxially puberulent, yellowbrown pubescent on veins or glabrescent. Bract of male flower 20–30 × 20–40 mm. Calyx segments lanceolate, apex acute.
Fruit ovoid or broadly ovoid, 5–7 cm, with longitudinal rows offlattened tubercles or undulate ridges, verrucose. Fl. Jun–Aug,
fr. Aug–Nov.
Roadsides on mountain slopes, thickets; 1400–2500 m. Yunnan[Bangladesh, India, Myanmar, Thailand].
The Chinese record of Momordica dioica Willdenow (FRPS
73(1): 192. 1986) is based on a misidentification of M. subangulata
subsp. renigera.

3. Momordica cochinchinensis (Loureiro) Sprengel, Syst. Veg.
3: 14. 1826.
木鳖子 mu bie zi
Muricia cochinchinensis Loureiro, Fl. Cochinch. 2: 596.
1790; Momordica macrophylla Gage; M. meloniflora HandelMazzetti; M. mixta Roxburgh.
Climbers, strong, to 15 m. Root tuberous. Stem andbranches glabrous or puberulent, sometimes tomentose at nodes.
Tendrils robust, simple. Petiole robust, 5–10 cm, slightly yel-low-brown pubescent or glabrescent, 2–4-glandular at base or
middle; leaf blade cordate or broadly ovate-orbicular, 10–20 ×
10–20 cm, 3–5-lobed; median lobe obovate or oblong-lanceolate, 6–10(–15) × 3–6(–9) cm, margin undulate-dentate, apex
acute or acuminate; lateral lobes ovate or oblong-lanceolate,
3–7(–11) × 2–4(–7) cm, base cordate. Plants dioecious. Male
flowers solitary or in a short raceme; pedicels robust, 3–5 cm,
or 6–12 cm when solitary, bracteate at apex; bract orbicularreniform, 3–5 × 5–8 cm, entire, retuse at apex; calyx tube funnelform; segments broadly lanceolate or oblong, 12–20 × 6–8
mm, apex acute or acuminate; corolla yellow; segments ovateoblong, 5–6 × 2–3 cm, yellow glandular at base, apex acute or
acuminate; stamens 3; anther cells reflexed. Female flower solitary; pedicel 5–10 cm, bracteate at middle; bract ca. 2 mm; calyx and corolla as in male flowers; ovary ovoid-oblong, ca. 1
cm, densely spinescent. Fruit red, ovoid, 12–15 cm in diam.,
fleshy, densely spinescent, apex rostellate. Seeds numerous, ashgray, ovoid or square, 26–28 × 18–20 mm, 5–6 mm thick, both
surfaces sculptured, margin undulate-sublobulate. Fl. Jun–Aug,
fr. Aug–Oct.
Forest margins and roadsides on mountain slopes; 400–1100 m.
Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hunan, Jiangsu, Jiangxi,
Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bangladesh, India, Malaysia, Myanmar].
The seeds, roots, and leaves are used medicinally.

13. ECBALLIUM A. Richard in Bory, Dict. Class. Hist. Nat. 6: 19. 1824, nom. cons.
喷瓜属 pen gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Elaterium Miller.
Herbs, creeping. Roots perennial. Stem and branches setose. Tendrils absent. Leaf blade cordate. Plants monoecious or dioecious. Male inflorescence a raceme; calyx tube shortly campanulate; segments 5, linear-lanceolate; corolla yellow, broadly campanulate or almost rotate; segments ovate-oblong, apex acute; stamens 3, linear; filaments short, free; anthers: one 1-celled, two 2-celled;
anther cells reflexed; connective broad, not produced; pistillode absent. Female flowers solitary; calyx and corolla as in male flowers;
staminodes 3, ligulate; ovary oblong, hispid; ovules numerous, horizontal; style short; stigmas 3, 2-lobed. Fruit oblong, hispid, scabrous, watery inside, when mature separating from peduncle and contracting at base, dispersing seeds by elastic contraction (squirting
seeds). Seeds numerous, brown or nearly black, oblong, compressed, narrowly marginate.
One species: Mediterranean region to SW Asia (east to Iran); introduced elsewhere, including China.

1. Ecballium elaterium (Linnaeus) A. Richard in Bory, Dict.
Class. Hist. Nat. 6: 19. 1824.
喷瓜 pen gua
Momordica elaterium Linnaeus, Sp. Pl. 2: 1010. 1753.
Roots elongate, robust. Stem ca. 1.5 m, scabrous. Petiole
somewhat robust, 5–15 cm, densely setose; leaf blade ovate-oblong or hastate, 8–20 × 6–15 cm, scabrous, abaxially white pubescent, adaxially white setose, margin sinuous or slightly divi-

ded, apex slightly obtuse; sinus semicircular, sometimes truncate. Male flowers in a raceme; peduncle densely yellow-brownpubescent and setose; pedicels filiform, 1–3 cm; calyx segments
lanceolate, 5–6 × 1.2–2 mm, densely pubescent and hispid; corolla yellow, ca. 3 cm in diam.; segments ovate-oblong; filaments ca. 3 mm, base villous. Fruit oblong or ovoid-oblong, 4–
5 × 1.5–2.5 cm, scabrous, yellow-brown hispid, both ends obtuse. Seeds ca. 4 mm.
Dry mountain slopes, grasslands. Xinjiang [native to SW Asia andMediterranean region].

14. SCHIZOPEPON Maximowicz, Mém. Acad. Imp. Sci. St.-Pétersbourg Divers
Savans 9 [Prim. Fl. Amur.]: 110. 1859.
裂瓜属 lie gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, scandent. Stem and branches slender. Tendrils 2-fid. Leaves long petiolate; leaf blade ovate-cordate or broadly ovate-

CUCURBITACEAE

31

cordate, rarely hastate, margin irregularly dentate, usually 5–7-lobed. Plants dioecious or monoecious; flowers small, unisexual or
rarely bisexual. Male flowers usually in a raceme; female flowers solitary or few in a raceme; calyx tube cupular or campanulate;segments 5, lanceolate or subulate; corolla segments 5, white, ovate; stamens 3, free or connate; filaments short; anthers: one 1celled, two 2-celled; anther cells erect; connective produced or not; ovary ovoid or conical, 3-locular or irregularly 3-locular, 1 ovulein each locule; ovule pendulous; style short, 3-fid or sometimes 5-fid; stigma slightly expanded, 2-lobed. Fruit small, ovoid or conical, smooth or punctate, apex acute or long acuminate, 3-valved or indehiscent, 1–3-seeded. Seeds pendulous, ovate, compressed,
margin irregularly dentate.
Eight species: E Asia to the Himalaya; eight species (five endemic) in China.

1a. Fruit conical, apex acuminate, rostrate; connective produced, subulate, rarely not produced; ovary irregularly
3-locular, ovule pendulous from middle of locule.
2a. Connective not produced ....................................................................................................................................... 6. S. bomiensis
2b. Connective produced, appendage subulate.
3a. Leaf blade ovate-cordate, usually unlobed, papery ...................................................................................... 7. S. bicirrhosus
3b. Leaf blade hastate, 5-lobed, membranous .................................................................................................... 8. S. xizangensis
1b. Fruit ovoid or ovate, apex acute; connective not produced; ovary 3-locular, ovule pendulous from apex of
locule.
4a. Flowers bisexual; stamens free .......................................................................................................................... 1. S. bryoniifolius
4b. Flowers unisexual; plant monoecious or dioecious; stamens connate.
5a. Plants monoecious; male and female flowers in same inflorescence, with many male flowers above
and 1 or 2 female flowers below; filaments connate, anthers free; ovary and fruit smooth ........................... 2. S. monoicus
5b. Plants dioecious.
6a. Filaments and anthers both connate; flowers larger; calyx segments ca. 3 mm; corolla segments
ca. 9 mm ................................................................................................................................................ 5. S. macranthus
6b. Filaments connate, anthers free or connate at base.
7a. Pedicels 2–5 mm; flowers small; calyx segments ca. 1 mm; corolla segments 2–3.5 mm; fruit
punctate ................................................................................................................................................... 3. S. dioicus
7b. Pedicels 5–10 mm; flowers slightly larger; calyx segments ca. 2 mm; corolla segments 3–5 mm .... 4. S. longipes
1. Schizopepon bryoniifolius Maximowicz, Mém. Acad. Imp.
Sci. St.-Pétersbourg Divers Savans 9 [Prim. Fl. Amur.]: 111.
1859.
裂瓜 lie gua
Schizopepon bryoniifolius var. japonicus Cogniaux; S. bryoniifolius var. paniculatus Komarov.
Plants annual. Stem 2–3 m. Branches slender, subglabrousor slightly puberulent. Tendrils filiform, glabrous. Petiole slender, 4–13 cm, sometimes puberulent; leaf blade ovate or
broadly ovate-cordate, 6–10 × 5–9 cm, membranous, puberulent, palmately 5–7-veined, 3–7-angular or -lobed, margin
irregularly dentate, apex acuminate; sinus semicircular. Flowers
very small, bisexual, solitary or in 3–5-flowered racemes; peduncle filiform, 1–1.5 cm, puberulent; calyx segments lanceolate, ca. 1.5 mm, 1-veined, entire; corolla rotate; segments white,ca. 2 × 0.8–1 mm, 3-veined; stamens inserted at base of calyx
tube, free, ca. 1 mm, glabrous; filaments linear; anthers ca. 0.5mm; anther cells erect, connective not produced; ovary ovoid,
3-locular; style short; stigmas 3. Fruit broadly ovoid, 10–15mm, apex acute, 3-valved, 1–3-seeded. Seeds ovate, ca. 9 × 5.5mm, compressed, margin irregularly dentate, apex truncate. Fl.
and fl. Jun–Nov.
River valleys, forests; 500–1500 m. Hebei, Heilongjiang, Jilin,
Liaoning [E Russia].

2. Schizopepon monoicus A. M. Lu & Zhi Y. Zhang, Acta
Phytotax. Sin. 23: 112. 1985.
峨眉裂瓜 e mei lie gua
Plants slender. Stem and branches glabrous, angular-sul-

cate. Tendrils filiform. Petiole slender, 3–4 cm, glabrous; leaf
blade ovate-cordate, 5–7 × 3–4 cm, membranous, palmately 5–
7-lobed; median lobe triangular, apex long acuminate; laterallobes acute, margin irregularly dentate, palmately 7-veined, both
surfaces setose at first; sinus semicircular. Plants monoecious.
Male and female flowers in same racemose inflorescence; peduncle slender, 2–4 cm, glabrous. Male flowers: pedicel filiform, 1–2 mm; calyx tube broadly campanulate; segments subulate, ca. 1.5 mm; corolla segments ovate, ca. 5 mm, 1-veined;filaments connate, ca. 0.3 mm; anthers free, ca. 0.4 mm; anthercells erect. Female flowers: pedicel ca. 2 mm; calyx segments
ovate, ca. 0.8 mm; ovary ovoid, ca. 0.8 mm, glabrous; style 3fid; stigma expanded. Fruiting pedicel ca. 1 cm, slender; fruit
subglobose, 3-locular with 1 seed in each locule. Fl. and fr.
May–Nov.
● Roadsides on mountain slopes. Sichuan (Emei Shan).

3. Schizopepon dioicus Cogniaux ex Oliver, Hooker’s Icon. Pl.
23: t. 2224. 1892.
湖北裂瓜 hu bei lie gua
Plants annual. Stem and branches slender. Tendrils filiform. Petiole 3.5–7.5 cm; leaf blade broadly ovate-cordate, 5–9
× 3–7 cm, membranous, usually 2- or 3-angled on each side,both surfaces glabrous, palmately 5–7-veined, margin dentate,apex acuminate; sinus semicircular. Plants dioecious. Male
flowers in a raceme; peduncle filiform, 5–10 cm; pedicels filiform, 2–3 mm; calyx segments linear-subulate or narrowly lanceolate, 1–1.2 × 0.5–0.8 mm; corolla white, rotate; segments
lanceolate or oblong-lanceolate, 2–3.5 × 1.5–2 mm, 1-veined;CUCURBITACEAE

32

filaments connate, ca. 0.8 mm; anthers free or united only atbase, ca. 0.5 mm; connective not produced. Female flowers
solitary or a few aggregated at apex of a short peduncle; ovary
ovoid, glabrous or pubescent, apex shortly acuminate, 3-locular,
1 ovule in each locule; styles 3-fid; stigma slightly expanded.
Fruiting pedicel filiform, 1–2 cm; fruit pale brown to blackbrown when mature, broadly ovoid, ca. 1.2 cm × 8 mm, glabrous or pubescent, smooth or verrucose, base rounded, apex
acute, 3-valved from apex, usually with 2 seeds. Seeds pale
brown when mature, ovoid, ca. 8 × 5 mm, base slightly wider,ca. 2 mm thick, irregularly dentate, contracted above, apex truncate.
● Grasslands, thickets, and roadsides on mountain slopes; 1000–
2400 m. Guizhou, W Hubei, NW Hunan, S Shaanxi, E and W Sichuan.

1a. Ovary and fruit densely pubescent ..... 3c. var. trichogynus
1b. Ovary and fruit glabrous.
2a. Leaf blade ovate-cordate, apex
acuminate ............................................ 3a. var. dioicus
2b. Leaf blade narrowly ovate-cordate,
apex caudate-acuminate ................... 3b. var. wilsonii
3a. Schizopepon dioicus var. dioicus
湖北裂瓜(原变种) hu bei lie gua (yuan bian zhong)Leaf blade ovate-cordate, apex acuminate. Ovary and fruit
glabrous; fruit pale brown when mature, verrucose.
● Grasslands and roadsides on mountain slopes; 1000–2400 m. W
Hubei, NW Hunan, S Shaanxi, E Sichuan.

3b. Schizopepon dioicus var. wilsonii (Gagnepain) A. M. Lu &Zhi Y. Zhang, Acta Phytotax. Sin. 23: 113. 1985.
四川裂瓜 si chuan lie gua
Schizopepon wilsonii Gagnepain, Bull. Mus. Natl. Hist.
Nat. 24: 378. 1918.
Leaf blade narrowly ovate-cordate, 3–5 lobed, apex caudate-acuminate. Ovary and fruit glabrous; fruit black-brown,smooth.
● Roadsides and thickets on mountain slopes; 1500–2400 m. NW
Guizhou, E and W Sichuan.

3c. Schizopepon dioicus var. trichogynus Handel-Mazzetti,
Oesterr. Bot. Z. 85: 219. 1936.
毛蕊裂瓜 mao rui lie gua
Ovary and fruit densely pubescent.
● Guizhou, W Hubei.

4. Schizopepon longipes Gagnepain, Bull. Mus. Natl. Hist.
Nat. 24: 378. 1918.
长柄裂瓜 chang bing lie gua
Stem and branches slender. Tendrils filiform. Petiole 4–7
cm, slightly puberulent; leaf blade ovate, 4–6 × 3–5 cm, irregularly 3–5-lobed; median lobe larger, triangular, apex acuminate; lateral lobes smaller, both surfaces puberulent, palmately
5-veined, margin dentate; sinus semicircular. Male flowers

many in a raceme, flowers on upper part of peduncle; rachis
filiform, 10–15 cm; pedicels linear, 0.5–1 cm, glabrous or puberulent; calyx segments lanceolate, 1.8–2 mm; corolla segments narrowly ovate, 3–5 × 1.5–2.5 mm, 1-veined, apex acuminate; filaments united, ca. 0.5 mm; anthers free, 0.8–1 mm;
connective not produced; cells erect, linear. Fl. Jul–Oct.
Valleys, roadsides; 2000–3000 m. W Sichuan, NW Yunnan [Bhutan, NE India (Darjeeling)].
This species was recorded from Bhutan and India in Fl. Bhutan(2(1): 260. 1991).

5. Schizopepon macranthus Handel-Mazzetti, Symb. Sin. 7:1064. 1936.
大花裂瓜 da hua lie gua
Plants annual. Stem and branches slender, sulcate-angular.
Tendrils filiform, glabrous. Petiole slender, 6–9 cm, subglabrous; leaf blade broadly ovate-cordate, 8–10 × 6–8 cm, membranous, both surfaces puberulent, palmately 7-veined, margin
irregularly dentate, apex caudate-acuminate; sinus subquadrate,
1–1.5 cm deep, 1.5–2 cm wide. Plants dioecious. Male flowers
in a raceme; peduncle slender, 6–8 cm; pedicels filiform, 2–5
cm; calyx segments lanceolate-subulate, ca. 3 mm; corolla paleyellow, rotate, glandular hairy at base; segments broadly lanceolate, ca. 9 mm, 1-veined, apex acuminate; filaments free, ca.
0.8 mm, glabrous; anthers connate at base, elliptic; cells erect,
puberulent; rudimentary ovary absent. Fl. Jul–Sep.
● Roadsides on mountain slopes; 2300–3000 m. W Sichuan, NW
Yunnan.

6. Schizopepon bomiensis A. M. Lu & Zhi Y. Zhang, Acta
Phytotax. Sin. 18: 385. 1980.
喙裂瓜 hui lie gua
Stems and branches slender, sulcate-angular, puberulent at
first or glabrescent. Tendrils slender. Petiole slender, 1–2(–3)
cm, subglabrous; leaf blade ovate-hastate or ovate-triangular,
2.5–5 × 2–3.5 cm, membranous, usually 5-lobed, lobe margindentate, both surfaces puberulent, apex shortly acuminate; sinus
suborbicular, 6–8 mm deep. Plants dioecious. Male flowers in a
raceme; peduncle pale yellow-green, 2.5–7 cm; pedicels ca. 3mm; calyx tube cupular; segments linear, ca. 1.2 mm; corollacampanulate; segments narrowly ovate, ca. 2 mm; filaments
nearly free, ca. 0.4 mm; anthers connate, one 1-celled, other 2celled, ca. 0.4 mm. Female flowers solitary in leaf axil; pedicels
filiform, 5–10 mm, glabrous; calyx segments subulate, 1.2–1.5
mm, apex acuminate; corolla yellow; segments narrowly ovate,ca. 2 × 1 mm, apex acuminate; staminodes glandular; ovary
conical, ca. 3 mm, puberulent, base rounded, apex elongate;style filiform. Fruiting pedicel slender, ca. 1 cm; fruit ca. 1.5 ×
0.5 cm, glabrous, base rounded, apex rostrate. Fl. and fr. Jul–
Oct.
● Forests or thickets on mountain slopes; 2200–2600 m. E and SE
Xizang.

7. Schizopepon bicirrhosus (C. B. Clarke) C. Jeffrey, Kew
Bull. 34: 802. 1980.
新裂瓜 xin lie gua

CUCURBITACEAE

33

Melothria bicirrhosa C. B. Clarke in J. D. Hooker, Fl. Brit.
India 2: 627. 1879; Schizopepon wardii Chakravarty.

glabrous; fruit narrowly conical, ca. 3 cm × 6 mm, white pubescent, apex attenuate, rostrate.

Branches ± slender, sulcate-angular, glabrous. Tendrils usually 2-fid, glabrous. Petiole slender, 2–4 cm, glabrous or puberulent; leaf blade 3–10 × 2–7 cm, membranous, both surfaces
slightly pubescent, margin undulate or 1- or 2-angled on eachside, irregularly dentate, apex caudate-acuminate; sinus semicircular. Plants dioecious. Male flowers 7–16 in a raceme; peduncle 3–7 cm; pedicels slender, 3–10 mm; calyx tube campanulate, 4–5 mm; segments subulate, ca. 2 mm, glandular
pubescent; corolla segments lanceolate, ca. 2.5 mm; stamensconnate, inserted on base of corolla tube; filaments ca. 1 mm,
glabrous; anthers puberulent; anther cells erect, ca. 1.5 mm;
connective produced; appendage ca. 1.5 mm. Female flowers
solitary; pedicels slender, 2–4 cm; calyx segments subulate, ca.
3 mm; corolla segments lanceolate, 5–6 mm, 3-veined; ovary
conical, ca. 8 × 3 mm, pubescent, base obtuse-rounded, apex
attenuate; ovules 3, solitary in each locule, pendulous; styles ca.
2 mm, 3-fid, apex expanded. Fruiting pedicel slender, 8–10 cm,

Thickets; 2700–2800 m. S Xizang [Bhutan, N India, Myanmar].

8. Schizopepon xizangensis A. M. Lu & Zhi Y. Zhang, Acta
Phytotax. Sin. 23: 116. 1985.
西藏裂瓜 xi zang lie gua
Plants glabrous. Branches slender, filiform. Tendrils filiform. Petiole slender, 2–3 cm; leaf blade hastate, 4–6.5 × 3–4
cm, thin, membranous, 5-lobed, lobe margin dentate, apex caudate-acuminate; sinus subquadrate. Plants dioecious. Male
flowers solitary, or 2–7 in a short raceme; calyx tube cupular,
ca. 3 mm in diam.; segments subulate, ca. 2 mm; corolla yellow; segments triangular-ovate, ca. 6 mm; stamens connate;filaments short, ca. 1 mm; anthers ovoid, one 1-celled, other 2celled; connective produced; appendage subulate, ca. 1 mm. Female flowers and fruit unknown.
● Herbage on mountain slopes; ca. 2100 m. SE Xizang (Mêdog).

15. BISWAREA Cogniaux, Bull. Soc. Roy. Bot. Belgique 21(2): 16. 1882.
三裂瓜属 san lie gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Warea C. B. Clarke, J. Linn. Soc., Bot. 15: 127. 1876, non Nuttall (1834).
Herbs, scandent. Tendrils 2-fid. Leaf blade 5–7-lobed. Plants dioecious; flowers yellow. Male flowers mostly racemose; calyx
tube narrowly tubular below, dilated above and broadly campanulate, 5-partite; segments linear; corolla broadly campanulate, 5partite; segments ovate, entire; stamens 3, inserted on calyx tube; filaments free; anthers connate, one 1-celled, others 2-celled; anthercells linear, triplicate; rudimentary ovary subulate. Female flowers: calyx and corolla as in male flowers; ovary oblong, with 3
placentas; styles filiform, elongate; stigmas 3, enlarged, ovules numerous, horizontal. Fruit oblong, 3-angled, 6-ribbed, fibrous, 3valved nearly to base. Seeds compressed, smooth.
One species: SW China, NE India, Myanmar, Nepal.

1. Biswarea tonglensis (C. B. Clarke) Cogniaux, Bull. Soc.
Roy. Bot. Belgique 21(2): 16. 1882.
三裂瓜 san lie gua
Warea tonglensis C. B. Clarke, J. Linn. Soc., Bot. 15: 129.
1876.
Stem and branches slender, puberulent or subglabrous.
Petiole robust, 6–10 cm; leaf blade broadly ovate-cordate or
suborbicular-cordate, 14–17 × 12–16 cm; lobes triangularovate, margin dentate, apex acute or acuminate. Male inflorescence usually 5–8-flowered; peduncle 8–10 cm; bracts 2–3 mm;calyx tube ca. 3 cm, base ca. 3 mm in diam., apex ca. 15 mm indiam.; segments narrowly lanceolate, 6–7(–20) mm, puberulent,
apex subulate; corolla segments ovate, 22–30 × ca. 11 mm;
apex acute; filaments filiform, 4–5 mm, glabrous; anthers connected, ca. 7 mm; rudimentary ovary subulate, 8–9 mm. Female
flower solitary; pedicel 25–30 mm; ovary densely tomentose,ca. 12 mm; calyx tube ca. 4.2 cm, apex ca. 11 mm in diam.;segments and corolla segments as in male flowers. Fruit 10–12
× 4–4.5 cm. Seeds numerous, pale brown, elliptic, 10–15 × 7–
10 mm.
NW Yunnan [NE India (Darjeeling, Sikkim), Myanmar, Nepal].

16. HERPETOSPERMUM Wallich ex J. D. Hooker in Bentham & J. D. Hooker,
Gen. Pl. 1: 834. 1867.
波棱瓜属 bo leng gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, climbing, annual. Roots spreading. Tendrils simple. Plants dioecious. Male flowers in raceme or rarely solitary; calyx
tube elongate, funnel-shaped; segments subulate; corolla broadly campanulate, 5-partite; segments entire; stamens 3, included incalyx tube; filaments free; anthers connate, one 1-celled, two 2-celled; anther cells linear, conduplicate; rudimentary ovary subulate.
Female flowers solitary; calyx and corolla as in male flowers; staminodes 3, subulate or absent; ovary oblong, 3-locular, 4–6 ovules

CUCURBITACEAE

34

in each locule, pendulous; filaments filiform, elongate; stigmas 3, dilated. Fruit broadly oblong, 3-valved from apex nearly to base.
Seeds pendulous, oblong or obovate, compressed, smooth, margins obtuse.
One species: Bhutan, SW China, India, Nepal.

1. Herpetospermum pedunculosum (Seringe) C. B. Clarke, J.
Linn. Soc., Bot. 15: 115. 1876.
波棱瓜 bo leng gua
Bryonia pedunculosa Seringe in Candolle, Prodr. 3: 306.
1828; Herpetospermum caudigerum Wallich ex Chakravarty;
H. grandiflorum Cogniaux.
Stem and branches slender, puberulent at first. Petiole 4–8(–10) cm; leaf blade ovate-cordate, 6–12 × 4–9 cm, membranous, both surfaces scabrous, yellow-brown pubescent at first,
base cordate, margin dentate, apex caudate-acuminate. Male

flowers usually in a raceme or solitary; inflorescence 12–40 cm,
5–10-flowered, puberulent; calyx tube 20–25 cm, apex 8–9 mm
in diam.; segments linear, 8–9 mm; corolla yellow; segments
elliptic, 20–22 × 12–15 mm; filaments 2–3 mm; anthers 5–6mm; rudimentary ovary subulate, 4–5 mm. Female flowers: calyx tube narrowly tubular, ca. 15 mm, densely yellow pubescent; segments linear, ca. 9 mm; corolla segments oblong, 20–
25 × 15–20 mm; ovary oblong-ovoid, densely yellow hispid.
Fruit 7–8 × 3–4 cm, fibrous. Seeds pale gray, oblong, ca. 12 × 5mm, 2–3 mm thick, base truncate. Fl. and fr. Jun–Oct.
Thickets and forest margins on mountain slopes; 2300–2500 m.
Xizang, Yunnan [Bhutan, India, Nepal].

17. EDGARIA C. B. Clarke, J. Linn. Soc., Bot. 15: 113. 1876.
三棱瓜属 san leng gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, climbing. Stem and branches slender. Tendrils 2-fid. Petiole slender; leaf blade ovate-cordate, membranous, pubescent at
first, ultimately glabrescent, scabrous, margin irregularly dentate. Plants dioecious. Male flowers in raceme or rarely solitary; calyx
tube elongate, narrowly funnelform; segments 5, linear; corolla yellow, subrotate, 5-partite; segments obovate, entire; stamens 3,
included, inserted on calyx tube; filaments free, short; anthers connate, one 1-celled, others 2-celled; anther cells straight; connective
narrow; pistillode subulate. Female flowers solitary; calyx and corolla as in male flowers; staminodes absent; ovary narrowly ovoid,
3-locular, 1 or 2 ovules in each locule; ovules ± horizontal; style filiform, elongate; stigmas 3, oblong, apex 2-lobed. Fruit broadly
fusiform, distinctly 3-angled, ribbed, dry, deeply 3-valved. Seeds subquadrate, compressed, slightly corrugate.
One species: Bhutan, SW China, N India, Nepal.

1. Edgaria darjeelingensis C. B. Clarke, J. Linn. Soc., Bot. 15:
114. 1876.
三棱瓜 san leng gua
Petiole 4–6 cm, pubescent at first, ultimately glabrous; leaf
blade ovate-cordate, 8–12 × 6–10 cm, membranous, abaxially
hispidulous on veins, adaxially sparsely puberulent at first, ultimately scabrous, margin sinuous-dentate, apex acute or shortly
acuminate; basal sinus 2–2.5 cm deep. Male flowers usually
10–16 in a raceme or rarely solitary; peduncle slender, 12–15

cm; pedicel filiform, 5–8 mm; calyx tube 1.5–2 cm, apex 3–4
mm in diam.; segments 3–4 mm; corolla segments narrowly
ovate, 10–15 × 3–5 mm, 3-veined, apex acuminate; filaments
ca. 5 mm; pistillode 3–4 mm. Fruiting pedicel stout, 2–10 cm;
fruit ovoid-3-angled, 6–7 cm, 3–4 cm in diam., glabrous orsparsely puberulent, apex 3-valved when mature. Seeds 13–14
× ca. 9 mm, ca. 2 mm thick. Fl. May–Aug.
Broad-leaved forests, grasslands; ca. 1700 m. Xizang [Bhutan, NIndia, Nepal].

18. LUFFA Miller, Gard. Dict. Abr., ed. 4, [806]. 1754.
丝瓜属 si gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, scandent, annual, glabrous or puberulent. Tendrils bi- or multifid. Petiole eglandular; leaf blade usually 5–7-lobed.
Plants monoecious. Male flowers in a raceme; calyx tube obconical; segments 5, triangular or lanceolate; corolla segments 5, free,spreading, entire or erose; stamens 3 or 5, free; anthers: one 1-celled, other 2-celled or all 1-celled when stamens 5; anther cells
linear, sigmoid-flexuous; connective usually dilated; pistillode glandular or absent. Female flowers solitary; calyx and corolla as inmale flowers; staminodes 3, or rarely 4 or 5; ovary cylindric; ovules numerous, horizontal; stigmas 3. Fruit oblong or cylindric, tereteor acutely costate, smooth or echinate, inside fibrous, apex operculately dehiscent. Seeds numerous, oblong, compressed.
Six species: tropical and subtropical regions throughout the world; two species (both introduced) in China.

1a. Stamens often 5, anther cells all 1-celled; fruit smooth, ecostate .............................................................................. 1. L. aegyptiaca
1b. Stamens 3, anthers: one 1-celled, others 2-celled; fruit with 8–10 acute ribs ........................................................... 2. L. acutangula

CUCURBITACEAE

35

1. Luffa aegyptiaca Miller, Gard. Dict., ed. 8, Luffa no. 1.
1768.

2. Luffa acutangula (Linnaeus) Roxburgh, Hort. Bengal. 70.
1814.

丝瓜 si gua广东丝瓜 guang dong si gua

Momordica luffa Linnaeus, Sp. Pl. 2: 1009. 1753; Luffa
cylindrica (Linnaeus) M. Roemer; M. cylindrica Linnaeus.
Stem and branches scabrous, sulcate-angular, puberulent.
Tendrils rather robust, usually 2–4-fid. Petiole 10–12 cm, scabrous, subglabrous; leaf blade triangular or suborbicular, 10–20
× 10–20 cm, often palmately 5–7-lobed; lobes triangular, median lobe 8–12 cm, base deeply cordate, margin dentate, apex
acute or acuminate; sinus 2–3 cm deep, 2–2.5 cm wide. Male
flowers usually 15–20 in a raceme; peduncle somewhat robust,
12–14 cm, pubescent; pedicels 1–2 cm; calyx broadly campanulate, 5–9 mm, puberulent; segments reflexed above, ovatelanceolate or subtriangular, 8–13 × 4–7 mm, densely puberulent, 3-veined, apex acuminate; corolla yellow, rotate, 5–9 cm
in diam.; segments oblong, 2–4 × 2–2.8 cm, inside densely yellow-white villous, base attenuate, apex rounded-obtuse. Stamens usually 5, rarely 3; filaments 6–8 mm, base white pubescent, connate at first, later free. Female flowers solitary;
pedicel 2–10 cm; ovary narrowly cylindric, puberulent; stigmas
expanded. Fruit cylindric, straight or slightly curved, 15–45 ×
3–6 cm, smooth, ecostate, inside strongly fibrous when mature.
Seeds usually black, ovate, smooth or very sparingly tuberculate, margin slightly winged. Fl. and fr. summer and autumn.
Commonly cultivated in China [widely cultivated throughout
tropical and temperate regions].

Cucumis acutangulus Linnaeus, Sp. Pl. 2: 1011. 1753;
Luffa acutangula var. subangulata (Miquel) Cogniaux; L. subangulata Miquel.
Stem rather robust, sulcate-angular, pubescent. Tendrils robust, often 3-fid, puberulent. Petiole 8–12 cm; leaf blade suborbicular, 15–20 × 15–20 cm, membranous, often 5–7-lobed;
median lobe broadly triangular, lateral lobes smaller, both surfaces puberulent, margin dentate, apex acute or acuminate;sinus subrounded. Male flowers 17–20 in raceme at apex ofpeduncle; peduncle 10–15 cm; pedicels 1–4 cm, white puberulent; calyx tube campanulate, 5–8 × ca. 10 mm; segments
lanceolate, 4–6 × 2–3 mm, densely white pubescent, 1-veined,
apex acuminate, slightly reflexed; corolla yellow, rotate; segments obcordate, 15–25 × 10–20 mm, both surfaces subglabrous; stamens 3, free; filaments 4–5 mm; anthers puberulent.
Female flowers solitary; ovary terete, longitudinally 8–10angled; style short; stigmas expanded, 2-lobed. Fruit cylindricor clavate-oblong, 8–10-angled, 15–40 × 6–10 cm, glabrous.
Seeds black, ovoid, 11–12 × 7–8 mm, ca. 1.5 mm thick, margin
without wing. Fl. and fr. summer and autumn.
Cultivated in S China [S and SW Asia; also cultivated in tropicalregions].
The fruit is eaten as a vegetable.

The fruit is eaten as a vegetable.

19. GYMNOPETALUM Arnott, Madras J. Lit. Sci. 12: 52. 1840.
金瓜属 jin gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Scotanthus Naudin; Tripodanthera M. Roemer.
Herbs, climbing, slender. Tendrils simple or 2-fid. Leaf blade ovate-cordate, angularly 3–5-lobed. Plants monoecious or dioecious. Male flowers solitary or in a raceme, ebracteate or bracteate; calyx tube elongate, tubular; segments 5, subulate; corolla whiteor yellow, rotate, 5-partite; segments oblong or obovate; stamens 3, inserted on middle of calyx tube; filaments short, free; anthers
connate, one 1-celled, others 2-celled; anther cells conduplicate; rudimentary ovary erect. Female flowers solitary; calyx and corolla
as in male flowers; staminodes 3, linear; ovary ovoid or oblong; ovules numerous, horizontal; placentae 3; style filiform. Fruit oblongor subglobose, acute or obtuse at both ends, indehiscent. Seeds obovate or oblong, compressed, marginate.
Four species: S Asia; two species in China.

1a. Leaf blade membranous, both surfaces slightly setose, 5-angular or 3–5-lobed or -partite, apex acuminate, margindentate; fruit oblong-ovoid, acute at both ends, 10-ribbed ............................................................................................ 1. G. chinense
1b. Leaf blade papery, both surfaces densely setose and villous, undivided or undulate 3–5-lobed, apex obtuse, margin
serrate; fruit subglobose, both ends obtuse, without ribs ............................................................................................... 2. G. scabrum
1. Gymnopetalum chinense (Loureiro) Merrill, Philipp. J. Sci.
15: 256. 1919.

florus (Roxburgh) Naudin, nom. superfl.; Tripodanthera cochinchinensis (Loureiro) M. Roemer.

金瓜 jin gua

Plants perennial. Stem and branches slender, hispid or
villous, glabrescent. Petiole 2–4 cm; leaf blade ovate-cordate,
4–8 × 4–8 cm, membranous, 5-angular or 3–5-lobed; middle
lobe larger, triangular, both surfaces scabrous, base cordate,
apex acuminate. Plants monoecious. Male flowers solitary, or
3–8 in a raceme; peduncle slender, 10–15 cm; bracts leaflike,
1–2.5 cm, yellow-brown villous, 3-lobed; calyx tube tubular,

Euonymus chinensis Loureiro, Fl. Cochinch. 1: 156. 1790;
Bryonia cochinchinensis Loureiro; Gymnopetalum cochinchinense (Loureiro) Kurz; G. heterophyllum Kurz; G. quinquelobatum Merrill; G. quinquelobum Miquel; Melothria touchanensis
H. Léveillé; Momordica tubiflora Roxburgh; Scotanthus tubi-

CUCURBITACEAE

36

elongate, ca. 2 cm; segments linear, ca. 7 mm; corolla white;segments oblong-ovate, 15–20 × 10–12 mm, ± villous; filaments ca. 0.5 mm; anthers ca. 7 mm. Female flowers solitary;
pedicels 1–4 cm; ovary oblong, 10–12 × ca. 5 mm, yellowbrown villous, acute at both ends; style 5–8 mm; stigmas 3.
Fruit orange, oblong-ovoid, 4–5 cm, smooth, 10-ribbed, acute atboth ends. Seeds oblong, ca. 7 × 3–3.5 mm, both ends obtuse.
Fl. Jul–Sep, fr. Sep–Dec.
Forests and thickets on mountain slopes; 400–900 m. Guangdong,
Guangxi, Hainan, Yunnan [Bhutan, India, Indonesia, Malaysia, Myanmar, Nepal, Philippines, Thailand, Vietnam].

2. Gymnopetalum scabrum (Loureiro) W. J. de Wilde &Duyfjes, Reinwardtia 12: 268. 2008.
风瓜 feng gua
Trichosanthes scabra Loureiro, Fl. Cochinch. 2: 589.
1790; Cucumis integrifolius Roxburgh; Gymnopetalum integrifolium (Roxburgh) Kurz; G. integrifolium var. penicaudii (Gagnepain) W. J. de Wilde & Duyfjes; G. leucostictum Miquel; G.
monoicum Gagnepain; G. monoicum var. incisum Gagnepain; G.
penicaudii Gagnepain; G. scabrum var. penicaudii (Gagnepain)

W. J. de Wilde & Duyfjes; Trichosanthes integrifolia (Roxburgh) Kurz.
Plants annual. Stem and branches slender, villous. Petiole
1.5–3 cm, densely villous; leaf blade reniform or ovate-cordate,
6–8 × 6–8 cm, papery, undivided or undulate 3–5-lobed; lobes
triangular, both surfaces scabrous, setose and villous, base cordate, margin serrate, apex obtuse. Plants monoecious. Male
flowers solitary, or in a raceme; peduncle 4–6 cm, densely villous; bracts leaflike, divided, densely yellow-brown villous;calyx tube 1.5–2 cm; segments lanceolate, ca. 8 mm; corollasegments obovate, 18–20 × 10–12 mm; filaments ca. 0.4 mm.
Female flowers solitary; pedicels 10–15 mm, densely villous;calyx and corolla as in male flowers; ovary ovoid-globose, ca.
10 mm, villous. Fruit orange-red when mature, subglobose, 2–3
cm in diam., smooth, not ribbed. Seeds narrowly oblong, ca. 9
× 3–3.5 mm, ca. 1.5 mm thick, both surfaces smooth, both ends
obtuse. Fl. Jun–Sep, fr. Sep–Nov.
Mountain slopes and herbage; 400–800 m. Guangdong, Guangxi,
Guizhou, Hainan, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia,
Myanmar, Philippines, Sri Lanka, Thailand, Vietnam].

20. HODGSONIA J. D. Hooker & Thomson, Proc. Linn. Soc. London 2: 257. 1854.
油渣果属 you zha guo shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Climbers, woody, large. Stem and branches stout. Tendrils robust, 2–5-fid. Plants dioecious. Male flowers in a raceme; calyx
tube elongate, campanulate; segments short; corolla rotate, 5-partite, lobes connate at base; segments obcuneate, apex truncate, very
long fimbriate; stamens 3; filaments inconspicuous; anthers connate, linear, one 1-celled, two 2-celled; anther cells conduplicate.
Female flowers solitary; calyx and corolla as in male flowers; ovary globose, 1-locular; ovules 12, horizontal; placentae 3, parietal
with pairs of ovules attached on each side; style long; stigma 3-lobed; lobes 2-fid, exserted. Fruit large, depressed, 12-grooved.
Perfect seeds usually 6, each having a rudimentary or barren seed attached to its side. Seeds large, flat, ellipsoid.
Two species: tropical Asia; one species in China.

1. Hodgsonia heteroclita (Roxburgh) J. D. Hooker & Thomson, Proc. Linn. Soc. London 2: 257. 1854.
油渣果 you zha guo
Trichosanthes heteroclita Roxburgh, Fl. Ind., ed. 1832, 3:705. 1832; Hodgsonia heteroclita subsp. indochinensis W. J. de
Wilde & Duyfjes.
Plants to 20–30 m. Stem and branches glabrous. Petiole
robust, 4–8 cm, striate, glabrous; leaf blade 15–24 × 15–24 cm,
leathery, both surfaces glabrous, 3–5-lobed, mostly 5-lobed;
lobes ovate-oblong, base truncate, apex acuminate. Male pe-duncle thick, 15–30 cm, striate, glabrous or puberulent; bracts
oblong-lanceolate, 5–10 mm, fleshy; pedicels short, thick, glabrous or puberulent; calyx tube yellowish, narrowly tubular, 7–10(–12) cm × 7–9 mm, dilated only at very apex; segments triangular-lanceolate, 2–4 mm; corolla yellow outside, white inside; segments ca. 5 cm, fimbriate fringes up to 15 cm. Female
pedicels robust, short; ovary subglobose, 2–2.2 cm in diam.
Fruit reddish brown, compressed globose, 10–16 × ca. 20 cm,
smooth or shallowly grooved. Seeds oblong, ca. 7 × 3 cm,
mostly compound. Fl. and fr. Jun–Oct.
Thickets, mountain slopes; 300–1500 m. Guangxi, SE Xizang, S
Yunnan [Bhutan, Cambodia, India, Laos, Myanmar, Thailand, Vietnam].

21. TRICHOSANTHES Linnaeus, Sp. Pl. 2: 1008. 1753.
栝楼属 gua lou shu
Huang Luqi (黄璐琦), Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Anguina Miller; Cucumeroides Gaertner; Involucraria Seringe.
Herbs, climbing, annual or perennial. Leaf blade simple, unlobed or palmately 3–7(–9)-lobed, rarely compound and 3–5-foliolate, margin usually denticulate. Tendrils usually 2–5-fid, rarely simple. Plants dioecious, rarely monoecious; flowers usually white,rarely pink or red. Male flowers usually in racemes, rarely solitary, sometimes male peduncles in axillary pairs, one 1-flowered,caducous, other bearing a raceme; bracts variable in size and form, rarely absent; calyx tube cylindric, frequently dilated at apex;

CUCURBITACEAE

37

segments 5, entire, serrate, or laciniate; corolla segments 5, usually long fimbriate; stamens 3, inserted on calyx tube; filaments very
short, free; anthers connate, two 2-celled, other 1-celled, cells conduplicate. Female flowers solitary, very rarely in racemes; calyx
and corolla as in male flowers; ovary inferior, ovoid or fusiform, 1-loculed with 3 parietal placentas; ovules usually many, generally
horizontal, half pendulous; style slender; stigmas 3, entire or bifid. Fruit globose, ovoid, or fusiform, fleshy, usually glabrous andsmooth, many seeded, indehiscent. Seeds packed in pulp, 1-loculed, oblong or ovate, and compressed, or 3-loculed, turgid, with 2
lateral locules empty.
About 100 species: Asia and N Australia; 33 species (14 endemic, one introduced) in China.

1a. Seeds subquadrangular, obovate-triangular, oblong, or 4-angled, 3-loculed, turgid, surrounded by thick ridges.
2a. Seeds 4-angled, with narrow middle ridge, 2 lateral locules broad; leaf blade abaxially glabrous or
puberulent on veins ................................................................................................................................... 33. T. tetragonosperma
2b. Seeds subquadrangular, obovate-triangular, or oblong, 2 lateral locules narrow; leaf blade abaxiallydensely pubescent.
3a. Leaves unlobed or irregularly shortly 3-lobed.
4a. Leaf blade ovate, thinly membranous, unlobed; fruit ovoid, smooth, glabrous ..................................... 30. T. baviensis
4b. Leaf blade broadly cordate, papery, unlobed or shortly 3-lobed; fruit ovoid-oblong or oblong,densely white villous ........................................................................................................................... 31. T. trichocarpa
3b. Leaves usually 3–7-lobed.
5a. Bracts lanceolate or oblanceolate, ca. 1.6 × 0.5–0.6 cm, margin triangular-dentate; leaves
adaxially glabrous or puberulent .................................................................................................................. 29. T. pilosa
5b. Bracts linear-lanceolate, 2–3 mm, entire; leaves adaxially tomentose ............................................ 32. T. cucumeroides
1b. Seeds elliptic, ovate-elliptic, or oblong, 1-loculed, compressed or turgid.
6a. Plants monoecious or dioecious; male flowers in racemes, bract very small, less than 5 mm; seed margin
undulate-dentate.
7a. Plants dioecious; bracts absent or ca. 5 mm; fruit cylindric, ca. 20 × 7 cm ................................................ 3. T. dafangensis
7b. Plants monoecious; bracts 1–3 mm or very small or absent; fruit cylindric and 100–200 cm, or
ovoid-oblong and 5–7 cm.
8a. Bracts 1–3 mm; fruit cylindric, 100–200 × 3–4 cm; leaf lobes obovate .................................................... 1. T. anguina
8b. Bracts absent or very small; fruit ovoid-oblong, 5–7 × 2.5–3.5 cm; leaf lobes rhombic ortriangular ................................................................................................................................................ 2. T. cucumerina
6b. Plants dioecious; male flowers solitary or in racemes or rarely in a panicle; bracts 1–6.5 cm or rarely
ca. 5 mm; seed margin crenellate.
9a. Leaves simple, leaf blade smooth on both surfaces; bract in male flower spreading, margin entire orsinuate-dentate.
10a. Male racemes densely villous.
11a. Leaves leathery, unlobed, leaf blade pubescent on veins ..................................................... 13. T. reticulinervis
11b. Leaves papery, unlobed or 3–5-lobed.
12a. Leaves unlobed; seeds subovate, 10–13 × 7–8 mm, margin undulate ..................................... 11. T. kerrii12b. Leaves 3–5-lobed or deeply lobed.
13a. Leaf blade abaxially densely white sericeous; male flowers solitary, densely
white sericeous ....................................................................................................... 12. T. sericeifolia
13b. Leaf blade abaxially densely brownish villous; male raceme densely brownish
villous ............................................................................................................................. 20. T. villosa
10b. Male racemes slightly hairy or villous, one peduncle 1-flowered, other bearing a raceme.
14a. Leaves ovate or ovate-oblong, subleathery or leathery, unlobed or rarely 3-lobed.
15a. Leaf blade papery, base cordate; fruit oblong, 7–10 × 6–7 cm ..................................... 10. T. homophylla
15b. Leaf blade subleathery or leathery.
16a. Leaf blade subleathery, unlobed, base rounded; fruit globose, ca. 7 cm
in diam. .................................................................................................................... 8. T. smilacifolia
16b. Leaf blade leathery, unlobed or shortly to deeply 2- or 3-lobed, basetruncate; fruit oblong, 12–18 × 5–10 cm ...................................................................... 9. T. truncata
14b. Leaves ovate-cordate or orbicular-cordate, papery, 3–9-lobed to middle or nearly to base.
17a. Male flowers solitary, smaller than 3 cm in diam.; calyx tube narrowly campanulate ..... 7. T. hylonoma
17b. Male flowers in raceme; flowers larger than 3 cm in diam.; calyx tube narrowly
funnelform or cylindric.
18a. Leaves usually 3–5(–7)-lobed.
19a. Bracts in male flowers ca. 3 cm, margin entire; calyx tube funnelform,ca. 3 cm, segments linear-lanceolate, 10–15 × 1.5–2.5 mm; leaves
ovate-cordate ......................................................................................... 4. T. jinggangshanica

CUCURBITACEAE

3819b. Bracts in male flowers less than 2.5 cm, margin sinuate-dentate;calyx tube cylindric, dilated at apex, 20–40 × ca. 10 mm; leaves
suborbicular ......................................................................................................... 5. T. kirilowii
18b. Leaves deeply 3–7-lobed usually up to base.
20a. Leaf lobes rhombic; fruiting peduncle 12–16.5 cm ................................. 6. T. mianyangensis
20b. Leaf lobes linear-lanceolate or oblanceolate; fruiting peduncle
4.5–8 cm ......................................................................................................... 14. T. rosthornii
9b. Leaves simple or 3–5-foliolate; leaf blade white punctate; bracts large, conspicuous, margin dentate.
21a. Leaves 3–5-foliolate; bracts rhombic-lanceolate.
22a. Leaflets remotely denticulate at margin .......................................................................................... 27. T. pedata22b. Leaflets entire at margin ......................................................................................................... 28. T. quinquefolia
21b. Leaves simple; bracts broadly ovate or oblong.
23a. Flowers red, pale red, or purple.
24a. Seeds ovoid, turgid ............................................................................................................. 22. T. dunniana
24b. Seeds oblong or oblong-ovate, compressed.
25a. Seeds oblong, truncate at both ends, ca. 18 mm; flowers purple; calyx tube
ca. 2.5 cm, segments linear; fruit orange-yellow ............................................... 25. T. rugatisemina
25b. Seeds oblong-ovate, 10–15 mm; flowers pale red.
26a. Leaf blade deeply (3–)5(–7)-lobed; calyx tube 4–6 cm, segments ovatelanceolate, denticulate; fruit red ....................................................................... 23. T. rubriflos
26b. Leaf blade shortly 5–7-lobed or lobed to middle; calyx tube ca. 4 cm,
segments narrowly lanceolate, entire; fruit orange-red ................................... 24. T. subrosea
23b. Flowers white.
27a. Bracts entire.
28a. Bracts ovate, 3.5–4.3 × ca. 2.5 cm, with 2 rows of brownish puncta
beside main vein; calyx segments 2–3 cm, margin sharply 2- or 3-lobed;
seeds triangular-ovate; leaves membranous, abaxially glabrous, adaxiallypunctate, petiole 5–11 cm ............................................................................... 18. T. quinquangulata
28b. Bracts oblong, ca. 4 × 1–1.5 cm; calyx segments ca. 1 cm, entire; seeds
subquadrangular; leaves papery, hirsute, petiole 14–17 cm ........................................ 19. T. cordata
27b. Bract margin dentate or lacerate.
29a. Leaves leathery.
30a. Bracts suborbicular, margin lacerate; calyx segments narrowly ovate,margin laciniate; leaves suborbicular .............................................................. 17. T. lepiniana
30b. Bracts obovate-oblong, margin dentate; calyx segments narrowly
lanceolate, margin dentate; leaves ovate-cordate ........................................ 26. T. tricuspidata
29b. Leaves membranous or papery.
31a. Leaves papery ............................................................................................ 21. T. laceribractea
31b. Leaves membranous.
32a. Leaf blade suborbicular, deeply palmately 5–7-lobed; bract margin
crenulate; calyx segments ca. 15 mm, entire ..................................... 15. T. wallichiana
32b. Leaf blade orbicular-cordate or broadly ovate-cordate, unlobed orirregularly 2- or 3(–5)-lobed; bract margin lobulate with linearlanceolate lobes; calyx segments ca. 10 mm, entire or serrate ....... 16. T. fissibracteata
1. Trichosanthes anguina Linnaeus, Sp. Pl. 2: 1008. 1753.
蛇瓜 she gua
Trichosanthes cucumerina Linnaeus var. anguina (Linnaeus) Haines.
Plants annual. Stems slender, branched, angular, puberulent. Petiole 3–7 cm; leaf blade suborbicular or orbicular-reniform, 8–16 × 6–18 cm, membranous, ± deeply 5(rarely 3–7)lobed; lobes usually obovate, abaxially densely puberulent,
adaxially subglabrous, margin finely denticulate. Plant monoecious. Male raceme usually 8–15-flowered; peduncle 10–18
cm, puberulent; pedicel erect, 5–12 cm, minutely spreading
bracteate, bracts 3(–5) mm; calyx tube subcylindric, slightly
dilated at apex, 2.5–3 × 4–5 mm, puberulent; segments re-flexed, ca. 2 mm. Female flowers solitary; peduncle up to 1 cm;
ovary narrowly fusiform, ± villous. Fruit very long, cylindric,
100–200 × 3–4 cm, smooth, sometimes contorted. Seed oblong,
11–17 × 8–10 mm, finely rugulose, margin undulate. Fl. and fr.
summer–autumn.
Cultivated in China [cultivated throughout the tropics].
The fruit is eaten as vegetable. The roots and seeds are used
medicinally for treating diarrhea.
One of us (Jeffrey) treats this taxon as Trichosanthes cucumerina
var. anguina.

2. Trichosanthes cucumerina Linnaeus, Sp. Pl. 2: 1008. 1753.
瓜叶栝楼 gua ye gua louCUCURBITACEAE

39

Trichosanthes brevibracteata Kundu; T. pachyrrhachis
Kundu.

Locally, the roots and fruit are used medicinally as “tian hua fen”
and “gua lou.”

Plants annual. Stems slender, profusely branched, ± pubescent. Leaf blade reniform or broadly ovate, (5–)7–10 × 8–11
cm, membranous, ± deeply 5–7-lobed; lobes triangular or
rhombic. Plants monoecious. Male peduncles in pairs, earlier 1flowered, later bearing a raceme; raceme few flowered; peduncle slender, 15–20 cm, puberulent; pedicel erect, 0.5–1.5 cm,
puberulent; bracts absent or very small; calyx tube somewhat
dilated at apex, 15–16 mm. Female flowers solitary or sometimes replacing earlier male flower; ovary oblong. Fruit ovoidoblong, 5–7 × 2.5–3.5 cm, with 7–10 seeds. Seeds ovate-oblong, 9–12 × 5–6 mm, compressed, rugulose, margin thick withtoothed projections from both surfaces. Fl and fr. autumn. 2n =22.

5. Trichosanthes kirilowii Maximowicz, Mém. Acad. Imp. Sci.
St.-Pétersbourg Divers Savans 9 [Prim. Fl. Amur.]: 482. 1859.

Forests or thickets along valleys, thickets on mountain slopes;
400–1600 m. Guangxi, S Yunnan [Bangladesh, India, Indonesia, Malaysia, Myanmar, Nepal, Pakistan, Sri Lanka; N Australia].
The roots, fruit, and seeds are used medicinally.
One of us (Jeffrey) treats this taxon as Trichosanthes cucumerina
var. cucumerina.

3. Trichosanthes dafangensis N. G. Ye & S. J. Li, Acta Phytotax. Sin. 27: 153. 1989.
大方油栝楼 da fang you gua lou
Leaf blade abaxially pale green, adaxially deep green, cordate, 10–22 × 8–20 cm, membranous, ± deeply palmately 5–7lobed; lobes elliptic-obovate, puberulent, margin distantly denticulate, apex acuminate. Male flowers solitary, fascicled, or inshort racemes; peduncle ca. 5 cm; bracts absent or ca. 5 mm;calyx tube funnelform, ca. 1 cm; segments linear-lanceolate, ca.
1 cm; pistillodes 3, claviform, ca. 5 mm. Female flowers solitary; pedicel ca. 2 cm; calyx tube shortly tubular; ovary ca. 2.5
cm, villous; staminodes 3. Fruit cylindric, ca. 20 × 7 cm, withca. 80 seeds. Seeds ovate, ca. 1 cm, compressed, verrucose,
margin sulcate, crenellate.
● Mountain slopes, thickets; 1000–1800 m. Guizhou.

4. Trichosanthes jinggangshanica C. H. Yueh, Acta Phytotax.
Sin. 18: 342. 1980.
井冈栝楼 jing gang gua lou
Stems slender, angular, puberulent. Petiole 3–8 cm, villous; leaf blade abaxially pale green, adaxially deep green, narrowly ovate-cordate to broadly ovate-cordate, 14–20 × 10–15
cm, papery, usually ± 3–5-lobed, rarely unlobed; middle lobe
ovate-triangular or ovate-lanceolate, abaxially glabrous or
villous on veins, adaxially villous. Male peduncles in axillarypairs, one 1-flowered, other bearing a raceme; peduncle 4–11
cm, villous; bracts ovate or orbicular-cordate, 2–2.5 × 1.5–4
cm, base rounded, margin entire, apex acuminate, shortly stipitate; calyx tube narrowly funnelform, ca. 3 cm; segments linearlanceolate, 10–15 × 1.5–2.5 mm. Female flowers solitary; pedicel ca. 1.5 cm; calyx tube ellipsoidal, ca. 2 cm; ovary oblong.
Fruiting peduncle ca. 6 cm; fruit oblong, 8–10 cm, smooth. Fl.
Mar–Jul, fr. Nov.
● Forests, thickets; 700–1500 m. SW Jiangxi.

栝楼 gua lou
Trichosanthes obtusiloba C. Y. Wu ex C. Y. Cheng & C.
H. Yueh.
Petiole 2–6 cm, striate, villous; leaf blade adaxially deep
green, suborbicular, 5–20 × 5–20 cm, papery, usually 3–5(–7)lobed up to middle, rarely deeply so (nearly up to base) or
unlobed; lobes rhombic-obovate or oblong, adaxially rough,margin sometimes remotely shortly lobed, apex obtuse. Male
raceme solitary or with solitary flowers in axillary pairs; peduncle 10–20 cm, robust, grooved, slightly hairy, 5–8-flowered atapex; bracts obovate or broadly ovate, 1.5–2.5(–3) × 1–2 cm,
margin sinuate-dentate, shortly puberulent, stipitate; calyx tube
cylindric, dilated at apex, 2–4 × ca. 1 cm; segments lanceolate,
10–15 × 3–5 mm, entire. Female flowers solitary; pedicel ca.
7.5 cm, shortly puberulent; calyx tube cylindric, ca. 2.5 × 1.2
cm; ovary oblong, 1.2–2 × ca. 1 cm. Fruiting peduncle 4–11
cm; fruit brown-yellow or orange-yellow, oblong or globose, 7–
10.5 cm. Seeds sandy beige, oblong-ovate, 11–16 × 7–12 mm,
marginate. Fl. May–Aug, fr. Aug–Oct. 2n = 88.
Open forests, shrublands, grasslands and fields beside villages;
200–1800 m. Gansu, Hebei, Henan, Jiangsu, Jiangxi, Shandong, Shanxi, Zhejiang [Japan, Korea].
This plant is the source of “tian hua fen” (the roots), “gua lou zi”(the seeds), and “gua lou pi” (the fruit rind), which are official source
drugs in traditional Chinese medicine.

6. Trichosanthes mianyangensis C. H. Yueh & R. G. Liao,
Bull. Bot. Res., Harbin 2(1): 115. 1992.
绵阳栝楼 mian yang gua lou
Stems densely shortly hairy and sparsely villous. Petiole
6–9 cm, punctate, puberulent; leaf blade orbicular-cordate, 11–
17 × 10–15 cm, papery, shortly 3–5-lobed or 7–9-lobed up to
middle; lobes rhombic, both surfaces shortly puberulent, margin
irregularly crenate-dentate, apex acuminate. Female flowers
solitary; pedicel 5–7 cm, slightly villous; calyx segments linear,
10–12 × 1–1.5 mm, entire; ovary oblong, 15–20 × 8–9 mm.
Fruiting peduncle 12–16.5 cm; fruit oblong, 10.5–11.5 × ca. 8.5
cm. Seeds up to 1.9 cm, smooth. Fl. May–Aug, fr. Aug–Oct. 2n= 88*.
● ?About 1000 m. Hubei, Sichuan.

7. Trichosanthes hylonoma Handel-Mazzetti, Symb. Sin. 7:1066. 1936.
湘桂栝楼 xiang gui gua lou
Trichosanthes leishanensis C. Y. Cheng & C. H. Yueh; T.
parviflora C. Y. Wu ex S. K. Chen.
Stems slender, with white lenticels. Petiole 3–9 cm; leaf
blade broadly ovate or broadly cordate, 11–17 × 10–16 cm, papery, usually 3–5-lobed up to middle, sometimes irregularly 5–
9-lobed; lobes oblong or rhombic-ovate, glabrous, margin re-CUCURBITACEAE

40

motely denticulate, apex acuminate. Male flowers solitary;
pedicel setiform, 4–7 cm; calyx tube narrowly campanulate,
1.2–1.5 × ca. 0.4 cm, glabrous or remotely villous; segments
erect or reflexed, linear, 6–7 mm. Female flowers unknown.
Fruit orange-red, ovoid-oblong, ca. 9 × 5–6 cm, attenuate atboth ends. Seeds brown, oblong, 1–1.3 × ca. 0.9 cm, baserounded, margin crenellate, apex truncate. Fl. May–Jun, fr.
Sep–Oct.
● Thickets along valleys; 800–1000 m. NE Guangxi, SE Guizhou,
S Hunan.

8. Trichosanthes smilacifolia C. Y. Wu ex C. H. Yueh & C. Y.
Cheng, Acta Phytotax. Sin. 18: 347. 1980.
菝葜叶栝楼 ba qia ye gua lou
Petiole 2–3 cm; leaf blade abaxially pale green, adaxiallydeep green, ovate-oblong, 9–13 × 7–9 cm, subleathery, unlobed,
both surfaces smooth, base rounded, margin reflexed, denticulate. Male flowers paniculate; peduncle 8–15 cm, slightly puberulent; bracts ovate-orbicular, ca. 1 × 0.8 cm, margin triangular-bidentate to middle, apex acuminate; calyx tube cylindric,ca. 2 × 0.5 cm; segments ca. 7 mm, long acuminate. Female
flowers unknown. Fruiting peduncle ca. 3 cm; fruit orange-yellow, globose, ca. 7 cm, smooth. Seeds brown, broadly oblong tosuborbicular, ca. 1.8 × 1.5 cm, both surfaces densely rugose,
margin undulate-crenate. Fl. autumn, fr. Nov–Dec.
● Evergreen broad-leaved forests; 600–1500 m. SE Xizang, S andSE Yunnan.

9. Trichosanthes truncata C. B. Clarke in J. D. Hooker, Fl.
Brit. India 2: 608. 1879.
大子栝楼 da zi gua lou
Trichosanthes crispisepala C. Y. Wu ex S. K. Chen; T.
ovata Cogniaux.
Stems robust, angular-striate, glabrous and punctate. Petiole 1.5–4(–6.5) cm; leaf blade ovate, narrowly ovate, or ovateoblong, 7–12 × 4.5–9 cm, leathery, unlobed or shortly to deeply
2- or 3-lobed, both surfaces glabrous, base rounded or truncate,margin usually entire or rarely distantly denticulate, apex acuminate. Male raceme 8–20-flowered; peduncle 7–15 cm, striate,flowering at apex; bracts ovate, shortly stipitate, margin entire
or more usually with a few teeth; calyx tube dilated at apex,
1.8–4 × ca. 1 cm, shortly tomentose; segments linear-lanceolate,
20–30 × 1–1.5 mm, entire. Female flowers solitary; ovary oblong, 1–2 × ca. 0.8 cm. Fruiting peduncle 4–5 cm; fruit oblong,
12–18 × 5–10 cm, glabrous. Seeds brown, oblong-obovate, 10–
23 × 5–20 mm, base truncate, apex obtuse. Fl. Apr–May, fr.
Jul–Aug. 2n = 22.
Montane forests, thickets; 300–1000 m. Guangdong, Guangxi, S
Yunnan [Bangladesh, Bhutan, India, Thailand, Vietnam].

10. Trichosanthes homophylla Hayata, Icon. Pl. Formosan.
10: 8. 1921.
芋叶栝楼 yu ye gua lou
Trichosanthes mushaensis Hayata.
Stems slender. Petiole 2–3 cm, hairy; leaf blade triangularcordate, 6–10 × 4–6 cm, papery, unlobed, margin dentate, adax-

ially scabrous, apex acuminate or acute. Male raceme 5–10flowered; peduncle 8–9 cm; pedicel ca. 3 mm; bracts obovate,
1–1.5 × 0.7–1.1 cm, margin irregularly crenate-dentate; calyx
tube cylindric, dilated at apex, ca. 3 × 0.8 mm; segments linear,ca. 6 mm. Female flowers unknown. Fruit oblong, 7–10 × 6–7
cm. Seeds elliptic, flattened. Fl. Aug.
Taiwan [Japan].

11. Trichosanthes kerrii Craib, Bull. Misc. Inform. Kew 1914:
7. 1914.
长果栝楼 chang guo gua lou
Trichosanthes tomentosa Chakravarty.
Stems robust, brownish hirsute. Petiole 2–5 cm, tomentose; leaf blade adaxially deep green, ovate-cordate, 10–20 × 7–
17 cm, papery, unlobed, abaxially densely felty-tomentose,
adaxially sparsely and shortly hirsute, margin entire or remotely
and obscurely serrate, apex long acuminate. Male peduncle ca.
2.5 cm; pedicel ca. 8 mm, tomentose; bracts linear, ca. 1.5 ×
0.25 cm; calyx tube cylindric, 2–3.5 × ca. 0.8 cm; segments
oblanceolate, ca. 1.8 × 0.45 cm, tomentose, apex acute. Female
peduncle robust, 2.5–3.5 cm, tomentose, 1-flowered; calyx tube
cylindric, ca. 1 cm, tomentose; segments narrowly triangular,
ca. 1 cm, apex long acuminate; ovary oblong, 1.5–2 × ca. 1 cm,
densely and softly tomentose. Fruiting peduncle robust, 2.5–5.5
cm, tomentose; fruit oblong, 8–10 × 4.5–6.5 cm. Seeds subovate, compressed, 10–13 × 7–8 mm, 1-veined at middle, baserounded or truncate, margin undulate and serrate, apex obtuse.
Forests along valleys or beside streams; 700–1900 m. SW Guangxi, SW Yunnan [India, Laos, Thailand, N Vietnam].

12. Trichosanthes sericeifolia C. Y. Cheng & C. H. Yueh, Acta
Phytotax. Sin. 18: 346. 1980.
丝毛栝楼 si mao gua lou
Stems angular-striate, sparsely sericeous. Petiole slender,
3.5–7 cm, white sericeous; leaf blade adaxially deep green,
ovate-cordate, 8–17 × 6–10 cm, papery, deeply 3–5-lobed;
middle lobes narrowly lanceolate or ovate-lanceolate, abaxiallydensely white sericeous, adaxially sparsely sericeous, margin
denticulate, apex acuminate. Male flowers solitary; pedicel
and flowering bud densely white sericeous; mature flowers
unknown. Female flowers solitary; pedicel 1.5–2.5 cm, densely
white sericeous; calyx tube tubular, ca. 1 × 0.4 cm, densely
white sericeous; segments linear-lanceolate, ca. 5 × 2 mm,
entire; ovary oblong, ca. 1.5 × 0.7 mm, densely white sericeous.
Fruit unknown. Fl. Apr–Jul.
● Thickets on mountain slopes or beside rivers; 700–1500 m. W
Guangxi, SW Guizhou, SE Yunnan.

13. Trichosanthes reticulinervis C. Y. Wu ex S. K. Chen, Bull.
Bot. Res., Harbin 5(2): 114. 1985.
两广栝楼 liang guang gua lou
Stems moderately robust, grooved, striate, glabrous or pubescent at nodes. Petiole 4–6 cm, striate, densely pubescent;leaf blade ovate to broadly ovate-cordate, 1.5–2 × 1–1.8 cm,
leathery, unlobed, both surfaces pubescent on veins, margin re-CUCURBITACEAE

motely denticulate, apex acuminate. Male flowers in racemes or
narrow panicles, 5–6 cm; pedicels ca. 6 mm, densely chocolatecolored pubescent; bracts lanceolate, ca. 1.5 cm, densely chocolate-colored villous; calyx tube campanulate, ca. 1.5 × 1.5 cm,
densely villous; segments triangular-ovate, ca. 1 × 0.6 cm.
Female flowers solitary; pedicel stout, ca. 2 cm, densely villous;calyx tube ca. 1 × 0.7 cm; segments linear, ca. 1.2 × 0.2–0.3 cm,
entire, densely villous; ovary ovoid, ca. 2.5 × 2 cm. Fruiting
peduncle ca. 2.5 cm, stout, densely villous; fruit ovoid, ca. 6 × 5
cm, densely villous. Seeds pale brown, ovate, compressed, ca.
1.1 × 0.7 cm, base rounded, apex obtuse. Fl. May–Jun, fr. Jul–Aug.
● Forests of low elevations; 200–400 m. Guangdong, Guangxi.

14. Trichosanthes rosthornii Harms in Engler, Bot. Jahrb.
Syst. 29: 603. 1901.
中华栝楼 zhong hua gua lou
Stems grooved, sparsely puberulent. Petiole 2.5–4 cm, striate, sparsely puberulent; leaf blade abaxially pale green, adaxially deep green, broadly ovate to suborbicular, (6–)8–12(–20)
× (5–)7–11(–16) cm, papery, deeply 3–7-lobed, usually deeply
5-lobed nearly to base; lobes linear-lanceolate, lanceolate, oroblanceolate, abaxially glabrous, adaxially shortly hairy, margin
denticulate, apex acuminate. Male raceme solitary or with one
1-flowered peduncle in axillary pairs; peduncle 8–10 cm; pedicels up to ca. 7 cm; bracts rhombic-obovate, 6–14 × 5–11mm, puberulent, attenuate at base, margin irregularly sinuatedentate; calyx tube narrowly cylindric, dilated at apex, 2.5–3(–3.5) cm. Female flowers solitary; pedicel 5–8 cm, puberulent; calyx tube cylindric, 20–25 × 5–8 mm; ovary oblong,
10–20 × 5–10 mm, puberulent. Fruiting peduncle 4.5–8 cm;
fruit orange-yellow, globose or oblong, 8–16 × 7–10 cm,
smooth, glabrous. Seeds brown, ovate-oblong, compressed,
1.5–2.2 × 0.8–1.4 cm, marginate. Fl. Jun–Aug, fr. Aug–Oct.
● Dense forests along valleys, thickets or grasslands on mountainslopes; 400–1900 m. S Anhui, N Guangdong, Guangxi, Guizhou, NE
Jiangxi, Sichuan, Yunnan.
The roots and fruit are used medicinally as “tian hua fen” and“gua lou.”

1a. Leaf blade deeply 5-lobed ......... 14d. var. huangshanensis
1b. Leaf blade deeply 3–7-lobed.
2a. Tendrils 4–6-fid ........................ 14c. var. multicirrata
2b. Tendrils 2- or 3-fid.
3a. Fruit 8–11 × 7–10 cm; seeds
1.5–1.8 × 0.8–0.9 cm ........... 14a. var. rosthornii
3b. Fruit 12–16 × 6.5–8.5 cm;
seeds 2–2.2 × 1.2–1.4 cm ..... 14b. var. scabrella

41● Guizhou, E Sichuan, Yunnan.

14b. Trichosanthes rosthornii var. scabrella (C. H. Yueh & D.
F. Gao) S. K. Chen, Fl. Reipubl. Popularis Sin. 73(1): 244. 1986.
糙籽栝楼 cao zi gua lou
Trichosanthes scabrella C. H. Yueh & D. F. Gao in C. H.
Yueh & C. Y. Cheng, Acta Phytotax. Sin. 18: 345. 1980.
Fruit large, 12–16 × 6.5–8.5 cm. Seeds 20–22 × 12–14mm, both surfaces slightly rugose.
● W Sichuan.

14c. Trichosanthes rosthornii var. multicirrata (C. Y. Cheng& C. H. Yueh) S. K. Chen, Fl. Reipubl. Popularis Sin. 73(1):244. 1986.
多卷须栝楼 duo juan xu gua lou
Trichosanthes multicirrata C. Y. Cheng & C. H. Yueh,
Acta Phytotax. Sin. 12: 430. 1974; T. damiaoshanensis C. Y.
Cheng & C. H. Yueh.
Leaf blade thick; lobes broad. Tendrils 4–6-fid, villous.
Calyx tube ca. 2 × 1.3 cm, densely pubescent.
● Forests, thickets, grasslands; 600–1500 m. N Guangdong,
Guangxi, Guizhou, Sichuan.

14d. Trichosanthes rosthornii var. huangshanensis S. K.
Chen, Bull. Bot. Res., Harbin 5(2): 116. 1985.
黄山栝楼 huang shan gua lou
Leaf blade deeply 5-lobed near to base; lobes further dissected; lobules 1 or 2, linear.
● S Anhui, NE Jiangxi.

15. Trichosanthes wallichiana (Seringe) Wight, Madras J. Lit.
Sci. 12: 52. 1840.
薄叶栝楼 bao ye gua lou
Involucraria wallichiana Seringe, Mém. Soc. Phys.
Genève 3(1): 25. 1825; Trichosanthes bracteata (Lamarck)
Voigt var. scotanthus (C. B. Clarke) Handel-Mazzetti; T.
grandibracteata Kurz; T. palmata Roxburgh var. scotanthus C.
B. Clarke.

Trichosanthes crenulata C. Y. Cheng & C. H. Yueh; T.
guizhouensis C. Y. Cheng & C. H. Yueh; T. stylopodifera C. Y.
Cheng & C. H. Yueh; T. uniflora K. S. Hao.

Stems branched, angular-grooved, glabrous. Petiole robust,
6–10 cm, striate, puberulent; leaf blade suborbicular, 18–20 ×
18–20 cm, membranous or papery, deeply (nearly to base) palmately 5–7-lobed; lobes oblong, abaxially glabrous or slightly
hairy on veins, adaxially glandular punctate and very rough,margin denticulate or irregularly crenate-dentate, apex acute or
acuminate. Male raceme 6–15-flowered at apex; peduncle stout,
10–20(–30) cm, grooved, puberulent; bracts broadly ovate, 2–3
cm, crenulate, slightly puberulent, multiveined; calyx tube ca. 5
× 1 cm; segments ca. 1.5 × 0.4 cm, entire. Female flowers
solitary; ovary ovoid-oblong, glabrous. Fruit ovoid or oblong,
5–10 × ca. 7 cm, apex acute or subacute. Seeds brown, oblong,
1.5–1.8 × 0.8–1.2 cm. Fl. Jul–Sep, fr. Oct–Nov.

Leaf blade deeply 3–7-lobed. Fruit 8–11 × 7–10 cm. Seeds
1.5–1.8 × 0.8–0.9 cm.

Forests along valleys; 900–2200 m. Xizang, S Yunnan [Bhutan,
India, Nepal].

14a. Trichosanthes rosthornii var. rosthornii中华栝楼(原变种) zhong hua gua lou (yuan bian zhong)CUCURBITACEAE

42

16. Trichosanthes fissibracteata C. Y. Wu ex C. Y. Cheng &C. H. Yueh, Acta Phytotax. Sin. 12: 438. 1974.
裂苞栝楼 lie bao gua lou
Petiole 6–10 cm, glabrous, striate; leaf blade adaxially palegreen, orbicular-cordate or broadly ovate-cordate, 11–25 × 9.5–
20 cm, membranous to papery, unlobed or irregularly shortly 2or 3-lobed or deeply 2- or 3-lobed, rarely 5-lobed; lobes broadly
triangular-ovate, rarely ovate-oblong, adaxially glabrous, withlarge disk glands at base, margin subentire or remotely minutely
denticulate, apex acuminate. Male raceme 9–20 cm; peduncle
glabrous or very shortly puberulent, 2–5-flowered at apex;
bracts ovate, 2.5–3 × 1.5–2 cm, cucullate, margin lobulate with
1.5–2 cm linear-lanceolate lobes; calyx tube ca. 3.5 cm; segments lanceolate, ca. 10 × 3–4 mm, entire or serrate. Female
flowers unknown. Fruit subglobose, ca. 6 cm. Seeds brown, oblong, 1.1–1.5 × 0.5–0.6 cm, base retuse, apex truncate or obtuse. Fl. Aug–Sep, fr. Nov.
● Dense forests along valleys, thickets on mountain slopes; 1100–
1300(–1500) m. SW Guangxi, SE Yunnan.

17. Trichosanthes lepiniana (Naudin) Cogniaux in A. Candolle & C. Candolle, Monogr. Phan. 3: 377. 1881.
马干铃栝楼 ma gan ling gua lou
Involucraria lepiniana Naudin in Huber, Cat. 11. 1868.
Stem robust, branched, glabrous. Petiole 4–7 cm, striate,glabrous or glandular-punctate; leaf blade adaxially deep green,
suborbicular, 9–17(–20) cm, shortly 3–5-lobed up to middle,
adaxially rough; lobe margin denticulate, apex acute or shortly
acuminate. Male raceme 13–17 cm; peduncle robust, striate,glabrous; pedicels ca. 5 mm; bracts suborbicular, ca. 4 cm, cucullate, margin lacerate; calyx tube attenuate from apex toward
base, ca. 7 cm, puberulent; segments narrowly ovate, ca. 1.5 ×
0.4 cm, margin laciniate. Female flowers solitary; pedicel 2.5–3
cm, glabrous, striate; bracts ovate, ca. 1.5 × 1 cm, entire; calyx
tube ca. 4 cm; ovary ovoid, ca. 1.5 × 1 cm, glabrous. Fruit red,
ovoid, ca. 9 cm in diam., smooth. Seeds brown, broadly ovate,ca. 1.5 × 0.8–1 cm, base obtuse or attenuate, apex truncate. Fl.
May–Jul, fr. Aug–Nov.
Evergreen broad-leaved forests along valleys, open forests andthickets on mountain slopes; 700–1200 m. SE Xizang, S and SE Yunnan[Bhutan, India].
De Wilde and Duyfjes (in Santisuk & Larsen, Fl. Thailand 9(4):520. 2008) considered the Chinese material here treated as Trichosanthes lepiniana to represent a distinct species, T. inthanonensis Duyfjes& Pruesapan (Thai Forest Bull., Bot. 32: 86. 2004, type from N Thailand; the type of T. lepiniana is from peninsular India).

18. Trichosanthes quinquangulata A. Gray, U.S. Expl. Exped.,
Phan. 645. 1854.
五角栝楼 wu jiao gua lou
Petiole 5–11 cm, striate, punctate; leaf blade adaxiallydeep green, quinquangular or broadly ovate, 13–22 × 12–20
cm, membranous, shortly 5-lobed or up to middle; lobes
broadly triangular or ovate-triangular, abaxially glabrous, adaxially glandular punctate and rough, margin denticulate, apex

long acuminate. Plants monoecious. Male raceme 17–30 cm;
peduncle glabrous, striate, 8–10-flowered; bracts ovate, 3.5–4.3
× ca. 2.5 cm, with 2 rows of brownish puncta beside main vein,
entire, apex long acuminate; calyx segments linear-lanceolate,
2–3 × 0.2–0.3 cm, margin sharply 2- or 3-lobed, apex longacuminate. Female flowers not seen. Fruiting peduncle 2–3 cm,
glabrous; fruit red, globose, 5–7 cm, glabrous. Seeds brown, triangular-ovate, 1–1.2 × 0.4–0.5 cm, base attenuate, apex triangular. Fl. Jun–Oct, fr. Sep–Oct.
Low-elevation thickets, roadsides; 500–900 m. Taiwan, S Yunnan[Indonesia, Malaysia, Myanmar, New Guinea, Philippines, Thailand,
Vietnam].

19. Trichosanthes cordata Roxburgh, Fl. Ind., ed. 1832, 3:703. 1832.
心叶栝楼 xin ye gua lou
Involucraria cordata (Roxburgh) M. Roemer; Trichosanthes microsiphon Kurz.
Stem robust, angular, grooved, glabrous or sparsely puberulent. Petiole 14–17 cm, striate, sparsely puberulent; leaf
blade broadly ovate-cordate, 8–22 × 7–23 cm, papery, unlobed
or rarely angular-sublobate, both surfaces scattered shortly hirsute, margin minutely denticulate, apex acute or shortly acuminate. Male raceme 4–8-flowered; peduncle 12–20 cm, stout,
striate, sparsely puberulent; pedicels 1–2 mm, thick; bracts oblong, ca. 4 × 1–1.5 cm, entire, shortly hirsute; calyx tube attenuate from apex toward base, ca. 4.5 × 0.7 cm; segments erect or
flexuous, linear-lanceolate, ca. 1.5 cm. Female flower solitary;
pedicel ca. 1 cm; ovary oblong, slightly puberulent. Fruit red,
globose, smooth. Seeds subquadrangular, 1–1.2 × 0.4–0.5 cm.
Fl. Jul–Aug, fr. Aug.
Evergreen broad-leaved forests; ca. 1000 m. SE Xizang [Bhutan,
India, Laos, Malaysia, Myanmar, Nepal, Singapore].

20. Trichosanthes villosa Blume, Bijdr. 934. 1826.
密毛栝楼 mi mao gua lou
Stems robust, long, densely brownish villous; petiole 10–
12 cm, densely brownish villous; leaf blade adaxially deep
green, broadly ovate, 11–18 × 11–17 cm, papery, unlobed or
tricuspidate, abaxially densely villous-hirsute, adaxially densely
and shortly brownish villous, margin denticulate, apex acuminate. Male raceme 10–20 cm, 15–20-flowered; peduncle
densely brownish villous; bracts oblong, 3–5 × 2–4 cm, both
surfaces brownish villous, margin irregularly dentate; calyx
tube 2.5–3 × 0.6–1.2 cm, densely brownish villous; segments
linear-lanceolate, ca. 2 × 0.3–0.4 cm, both surfaces villous,
margin entire, apex long acuminate. Female flowers solitary;
pedicel ca. 1.5 cm, densely long villous; calyx tube cylindric,ca. 1.5 × 0.7 cm; ovary oblong, ca. 1.5 × 1 cm, densely villous.
Fruit brown-red, subglobose, 8–13 cm. Seeds oblong or obovate-triangular, 1.7–2.8 × 1–1.7 cm, 1-veined at middle, baseobtuse, apex truncate. Fl. Dec–next Jul, fr. Sep–Nov.
Thickets or open forests on mountain slopes; 300–1000 m. S
and SW Guangxi, S Yunnan [Indonesia, Laos, Malaysia, Philippines,
Vietnam].

CUCURBITACEAE

21. Trichosanthes laceribractea Hayata, J. Coll. Sci. Imp.
Univ. Tokyo 30(1): 117. 1911.
长萼栝楼 chang e gua lou
Trichosanthes hupehensis C. Y. Cheng & C. H. Yueh; T.
koshunensis Hayata; T. punctata Hayata; T. schizostroma
Hayata; T. shikokiana Makino; T. sinopunctata C. Y. Cheng &C. H. Yueh.
Stems angular, grooved, subglabrous. Petiole 1.5–9 cm,
striate, hispidulous or white punctate; leaf blade abaxially palegreen, adaxially deep green, suborbicular or broadly ovate, 5–16(–19) × 4–15(–18) cm, papery, 3–7-lobed shortly or nearly tobase; lobes triangular, ovate, or rhombic-obovate, abaxially hispidulous on veins, adaxially densely hispidulous or white punctate, margin remotely mucronate-serrate, apex acuminate. Malepeduncle robust, 10–23 cm, puberulent or sparsely hispidulous;
bracts broadly ovate, 2.5–4 × 2.5–4 cm, cucullate, margin
lacerate; calyx tube narrowly tubular, ca. 5 cm; segments erect,
ovate, 1–1.3 × ca. 0.7 cm, margin laciniate, apex acuminate.
Female flowers solitary; pedicel 1.5–2 cm, puberulent; calyx
tube cylindric, ca. 4 × 5 cm; segments linear, 1–1.3 cm, entire;
ovary ovoid, ca. 1 × 0.7 cm, smooth. Fruit globose to ovoidglobose, 5–8 cm in diam. Seeds oblong, 1–1.4 × 0.5–0.8 cm,
both ends rounded or truncate. Fl. Jul–Aug, fr. Sep–Oct. 2n =22*.
● Dense forests along valleys; 200–1100 m. Guangdong, Guangxi, Hubei, Jiangxi, Sichuan, Taiwan.

22. Trichosanthes dunniana H. Léveillé, Repert. Spec. Nov.
Regni Veg. 10: 148. 1911.
糙点栝楼 cao dian gua lou
Trichosanthes prazeri Kundu; T. rubriflos Thorel ex Cayla
f. macrosperma C. Y. Cheng & C. H. Yueh; T. tridentata C. Y.
Cheng & C. H. Yueh.
Stem moderately robust, grooved, punctate. Petiole slender, 4–5 cm, punctate; leaf blade abaxially pale green, adaxiallygreen, suborbicular, 10–15 cm, papery, deeply palmately 5–7lobed; lobes elliptic-obovate, abaxially glabrous, glandular
punctate on veins, adaxially glandular punctate, narrowed
toward base, cuspidate. Male racemes 8–10 cm; peduncle stout,
striate, densely pubescent or punctate, 5–10-flowered; bracts
brown, broadly ovate, ca. 5 × 4 cm, abaxially punctate, margin
deeply serrate; calyx tube ca. 5 cm; segments pale red. Female
flowers unknown. Fruit oblong, ca. 8 × 6 cm, smooth, apex
acute. Seeds ovoid, turgid, ca. 1.2 × 0.8 cm. Fl. Jul–Sep, fr.
Oct–Nov.
Dense forests along valleys or thickets; 900–1900 m. Guangxi,
Guizhou, Sichuan, Yunnan [Laos, Myanmar, N Thailand].

23. Trichosanthes rubriflos Thorel ex Cayla, Bull. Mus. Natl.
Hist. Nat. 14: 170. 1908.
红花栝楼 hong hua gua lou
Trichosanthes majuscula (C. B. Clarke) Kundu; T. multiloba Miquel var. majuscula C. B. Clarke; T. pubera Blume
subsp. rubriflos (Thorel ex Cayla) Duyfjes & Pruesapan.

43

Stems robust, branched, grooved, pubescent. Petiole 5–12(–18) cm, striate, pubescent; leaf blade adaxially deep green,broadly ovate or suborbicular, 7–20 cm, deeply palmately 3–7lobed; lobes broadly ovate, oblong or lanceolate, abaxiallypubescent, adaxially hispidulous or glandular punctate, margin
denticulate or irregularly serrate, apex acuminate. Male raceme
10–20 cm; peduncle grooved, puberulent, (6–)11–14-flowered;bracts red, broadly ovate or obovate-rhombic, 2.5–4 × ca. 3 cm,
puberulent, margin lacerate; calyx tube red, 4–6 cm; segments
linear-lanceolate, 1.2–1.6 × 0.3–0.5 cm, margin entire or rarely
denticulate, apex acuminate; corolla red or pale red. Female
flowers solitary; pedicel ca. 2 cm, densely pubescent; calyx
tube ca. 3 cm; ovary ovoid, ca. 2 × 1 cm. Fruiting peduncle robust, ca. 3 cm, striate, pubescent; fruit red, broadly ovoid or
globose, 7–9.5 × 5.5–8 cm, smooth, apex acute. Seeds yellowbrown, oblong, ca. 1.2 × 0.4–0.5 cm, base truncate, apex attenuate. Fl. May–Nov, fr. Aug–Dec.
Dense forests along valleys, open forests or thickets on mountainslopes; (100–)400–1600 m. Guangdong, Guangxi, Guizhou, Xizang,
Yunnan [Cambodia, NE India, Laos, Myanmar, Thailand, Vietnam].

24. Trichosanthes subrosea C. Y. Cheng & C. H. Yueh, Acta
Phytotax. Sin. 18: 349. 1980.
粉花栝楼 fen hua gua lou
Stem glabrous, grooved. Petiole 4–6.5(–8) cm, striate, glabrous; leaf blade abaxially pale green, adaxially deep green,quinquangular-cordate or suborbicular, 11–15 cm, thinly papery, shortly 5–7-lobed or to middle; central lobe triangular or
subrhombic, abaxially glabrous, adaxially hispidulous or glandular-punctate, margin triangular-dentate, apex long acute. Malepeduncles occasionally in pairs, one early 1-flowered and otherbearing a raceme; pedicel ca. 8 cm, striate, glabrous; bract
ovate, 2.5–3 cm, cucullate, glabrous, subentire, apex obtuse; calyx segments narrowly lanceolate, 1.5–2 cm, entire; corolla pale
red. Female flowers unknown. Fruiting peduncle stout, 2–2.5
cm; fruit orange-red, subglobose, 7–9 × 5–7.5 cm, smooth.
Seeds oblong, base truncate, apex triangular. Fl. Jul–Aug, fr.
Aug–Sep.
Forests; ca. 1700 m. S Guangxi, SW Yunnan [Myanmar].

25. Trichosanthes rugatisemina C. Y. Cheng & C. H. Yueh,
Acta Phytotax. Sin. 12: 440. 1974.
皱籽栝楼 zhou zi gua lou
Stem grooved, densely villous. Petiole 3.5–5.5 cm, densely
villous; leaf blade suborbicular, 10–15 cm, papery, 3–5-lobed
up to middle; lobes ovate or obovate-elliptic, both surfaces pubescent, margin triangular-denticulate, apex acuminate. Male
racemes 3–6 cm, densely brown villous, 3–5-flowered at apex;
bracts ovate, 2.5–3.5 × 2–3 cm, cucullate, apex acuminate; calyx tube ca. 2.5 cm; segments ovate, ca. 15 mm, margin dentate;corolla red. Female flowers solitary; calyx tube cylindric, ca. 3
cm; segments linear, pubescent; ovary oblong, ca. 1.5 × 0.5 cm.
Fruiting peduncle ca. 2 cm; fruit orange-yellow, ovoid, 8–9.5 ×
5–6 cm, attenuate at apex. Seeds gray-green, ovate-oblong, ca.
1.5 × 0.7 cm, both ends truncate.
● Thickets. SE and SW Yunnan.

44

CUCURBITACEAE

26. Trichosanthes tricuspidata Loureiro, Fl. Cochinch. 2: 589.
1790.
三尖栝楼 san jian gua lou
Modecca bracteata Lamarck; Trichosanthes bracteata(Lamarck) Voigt.
Stems robust, branched, angular-striate, grooved, glabrous.
Petiole ca. 5 cm, striate, glabrous, punctate; leaf blade adaxiallydeep green, broadly ovate-cordate, 12–13 × 12–13 cm, thinly
leathery, shortly 3-lobed; lobes ovate-triangular, abaxially glabrous, adaxially hispidulous or punctate, margin undulate-denticulate, apex acute or shortly acuminate. Male raceme 12–15
cm; peduncle stout, striate and grooved, glabrous; pedicel short;bracts obovate-oblong, ca. 4 × 2–2.5 cm, cucullate, both surfaces white squamose or pubescent, margin dentate, apex obtuse; calyx tube narrowly funnelform, ca. 6 × 1 cm; segments
narrowly lanceolate, ca. 1 × 0.3 cm. Female flowers and fruit
not seen.
Thickets on mountain slopes; ca. 900 m. Guizhou [Indonesia,
Malaysia, Nepal, Thailand, Vietnam].

27. Trichosanthes pedata Merrill & Chun, Sunyatsenia 2: 20.
1934.
趾叶栝楼 zhi ye gua lou
Trichosanthes pedata var. yunnanensis C. Y. Cheng & C.
H. Yueh.
Stem slender, striate, grooved, glabrous or pubescent at
nodes. Leaves 3–5-foliolate; petiole 2.5–6 cm; leaflets abaxiallypale green, lanceolate or oblong-oblanceolate, 9–12 × 2.5–3.5
cm, membranous, abaxially glabrous, adaxially hispidulous or
white punctate, remotely denticulate, acuminate; petiolule 2–5(–11) mm. Male raceme 14–19 cm; peduncle and pedicels
brown pubescent; bracts obovate or rhombic-ovate, 1–1.5 × ca.
0.8 cm, pubescent, margin entire or dentate, apex acuminate.
Female flowers solitary; calyx tube cylindric, ca. 3 × 5 cm;
ovary ovoid, ca. 1.5 × 0.8 cm, glabrous. Fruiting peduncle 1(–3)
cm; fruit orange-yellow, globose, 5–6 cm, smooth. Seeds brown,
ovoid, tumid, 1–1.2 × ca. 0.8 cm, base triangular, apex rounded.
Fl. Jun–Aug, fr. Jul–Dec. 2n = 22.

(3–)5–8 × ca. 3 mm, pubescent, margin entire, apex acuminate.
Female flowers unknown. Fruit oblong, ca. 6 × 4 cm, smooth,
apex acute. Seeds brown, ovoid-oblong, tumid, 12–15 × 8–10mm, base rounded. Fl. Aug–Sep, fr. Oct–Nov.
Open forests; 900–1400 m. S Yunnan [Laos].

29. Trichosanthes pilosa Loureiro, Fl. Cochinch. 2: 588. 1790.
全缘栝楼 quan yuan gua lou
Trichosanthes chingiana Handel-Mazzetti; T. himalensis
C. B. Clarke; T. himalensis var. indivisa Chakravarty; T. okamotoi Kitamura; T. ovigera Blume; T. ovigera var. sikkimensis
Kundu.
Stems slender, grooved, pubescent. Petiole 4–12 cm, striate, densely pubescent; leaf blade ovate-cordate to suborbicular,
7–19 × 7–8 cm, papery, unlobed or very shortly 3-lobed or 3–5lobed to middle or to base; median lobe ovate, oblong, or obovate-oblong, abaxially pubescent, adaxially glabrous or very
shortly puberulent, margin distantly denticulate or dentate, apex
acuminate. Male peduncle 10–26 cm, striate, densely pubescent; bracts lanceolate or oblanceolate, ca. 1.6 × 0.5–0.6 cm,
attenuate to base, margin triangular-dentate, apex acuminate;calyx tube subcylindric, slightly dilated at apex, ca. 5 cm;
segments triangular-ovate, 7–10 × 2–3 mm, entire. Female
flowers solitary; pedicel 1–1.3 cm, striate, densely pubescent;calyx tube cylindric, 2–5 × 0.2–0.3 cm, glabrous or pubescent;ovary ovoid, 1–1.5 × 0.3–0.5 cm. Fruit ovoid or oblong, 5–7 ×
2.5–4 cm, smooth, base rounded, apex acuminate, beaked. Seeds
subquadrangular, 7–9 × 7–8 mm, longitudinally surrounded by
2 thick ridges, thick near middle; 3-loculed with 2 lateral locules empty. Fl. May–Sep, fr. Sep–Dec. 2n = 22.
Thickets along valleys, open forests on mountain slopes; 700–
2500 m. Guangdong, Guangxi, Guizhou, Yunnan [India, Indonesia,
Japan, Nepal, Thailand, Vietnam].
One of us (Jeffrey) treats the following three species (i.e., Trichosanthes baviensis, T. trichocarpa, and T. cucumeroides) as synonyms
of T. pilosa.

30. Trichosanthes baviensis Gagnepain, Bull. Mus. Natl. Hist.
Nat. 24: 379. 1918.

Open forests along valleys, thickets; 200–1500 m. Guangdong,
Guangxi, Hainan, Hunan, Jiangxi, Yunnan [Vietnam].

短序栝楼 duan xu gua lou

28. Trichosanthes quinquefolia C. Y. Wu ex C. Y. Cheng & C.
H. Yueh, Acta Phytotax. Sin. 18: 351. 1980.

Stems slender, grooved, glabrous or pubescent. Petiole
slender, 4–9 cm, striate, densely pubescent; leaf blade abaxiallypale green, adaxially deep green, ovate, 5–20 × 5–13 cm, thinly
membranous, unlobed, abaxially pubescent, adaxially puberulent, margin denticulate, apex acuminate. Male flowers in corymb, ca. 2 cm; pedicel 1–1.5 cm, pubescent; bracts absent; calyx tube dilated at apex, ca. 2 cm, pubescent; segments narrowly triangular, ca. 3 mm. Female flowers solitary; pedicel ca.
7 mm, pubescent; ovary oblong, 8–10 mm, densely pubescent.
Fruit ovoid, 3.5–5 × ca. 3.5 cm, smooth, glabrous, beaked.
Seeds unknown. Fl. Apr–May, fr. May–Sep.

木基栝楼 mu ji gua lou
Stems striate, glabrous or pubescent at nodes. Petiole 3–4
cm, pubescent; leaf blade 3–5-foliolate; leaflets sessile; medianleaflet abaxially pale green, lanceolate, 15–23 × 2–4 cm, membranous or thinly papery, abaxially glabrous, adaxially hispidulous or white punctate, base cuneate, margin entire, apex acuminate. Male raceme 11–26 cm; peduncle striate, pubescent, 5–15-flowered; bracts rhombic, 7–10 × ca. 5 mm, pubescent,margin crenate, apex obtuse; calyx tube narrowly funnelform,ca. 3 × 0.7 cm, densely pubescent; segments linear-lanceolate,● Evergreen broad-leaved forests or thickets; 600–1500 m.
Guangxi, SW Guizhou, S and SE Yunnan.

CUCURBITACEAE

One of us (Jeffrey) treats this taxon as a synonym of Trichosanthes pilosa.

31. Trichosanthes trichocarpa C. Y. Wu ex C. Y. Cheng & C.
H. Yueh, Acta Phytotax. Sin. 18: 340. 1980.
杏籽栝楼 xing zi gua lou
Stems slender, grooved, puberulent. Petiole slender, 7–8
cm, striate, puberulent; leaf blade abaxially pale green, adaxially deep green, broadly cordate, 10–16 × 7–13 cm, papery,
unlobed or irregularly shortly 3-lobed; lobes triangular, abaxially pubescent, adaxially sparsely hairy, margin entire or denticulate, apex acuminate to long acuminate. Flowers unknown.
Fruiting peduncle 3–4 cm, striate, puberulent; fruit yellow ororange-red, ovoid-oblong or oblong, 7–9 × 7–8 cm, densely
white villous, beaked. Seeds obovate-triangular, 1–1.1 × ca. 1
cm, 3-loculed with 2 lateral locules narrow. Fr. Oct–Nov.
● Forests on mountain slopes, roadsides; 2100–2400 m. S and SW
Yunnan.
One of us (Jeffrey) treats this taxon as a synonym of Trichosanthes pilosa.

45

王瓜(原变种) wang gua (yuan bian zhong)
Bryonia cucumeroides Seringe in Candolle, Prodr. 3:308. 1828; Trichosanthes cavaleriei H. Léveillé; T. chinensis
Seringe; T. cucumeroides var. formosana (Hayata) Kitamura;T. formosana Hayata; T. ovigera Blume subsp. cucumeroides(Seringe) C. Jeffrey.
Fruit ovoid or ovoid-oblong. Seeds oblong, middle ridge
as broad as 2 lateral locules.
Guangdong, Guangxi, Jiangxi, Sichuan, Taiwan, Zhejiang [Japan].

32b. Trichosanthes cucumeroides var. hainanensis (Hayata)
S. K. Chen, Bull. Bot. Res., Harbin 5(2): 117. 1985.
海南栝楼 hai nan gua lou
Trichosanthes hainanensis Hayata, Icon. Pl. Formosan. 10:
8. 1921.
Fruit ovoid or ovoid-oblong. Seeds triangular-ovate, withmiddle ridge broad; 2 lateral locules narrow, ca. 2.5 mm indiam.
● Guangdong, Guangxi, Hainan.

32. Trichosanthes cucumeroides (Seringe) Maximowicz inFranchet & Savatier, Enum. Pl. Jap. 1: 172. 1873.

32c. Trichosanthes cucumeroides var. stenocarpa Honda,
Bot. Mag. (Tokyo) 54: 223. 1941.

王瓜 wang gua

狭果草栝楼 xia guo cao gua lou

Stems slender, branched, grooved, pubescent. Petiole 3–10
cm, striate, densely pubescent; leaf blade abaxially pale green,
adaxially deep green, broadly ovate or orbicular, 5–13(–19) ×
5–12(–18) cm, papery, usually shortly 3–5-lobed, sometimes
unlobed; lobes triangular or ovate to obovate-oblong, abaxiallydensely shortly pubescent, adaxially tomentose, margin denticulate or serrate, apex obtuse or acuminate. Male peduncle 5–
10 cm, striate, pubescent; bracts linear-lanceolate, 2–3 mm, pubescent, entire; calyx tube 6–7 cm, pubescent; segments linearlanceolate, 3–6 × ca. 1.5 mm, entire, acuminate. Female flowers
solitary; pedicel 0.5–1 cm; ovary oblong, densely pubescent.
Fruiting peduncle 5–20 mm, pubescent; fruit orange-red, ovoid,
ovoid-oblong, or globose, 6–7 × 4–5.5 cm, smooth, beaked.
Seeds deep brown, oblong or triangular-ovate, 7–12 × 7–14
mm, 3-loculed; 2 lateral ones large, ca. 4.5 mm in diam. Fl.
May–Aug, fr. Aug–Nov.
Dense forests along valleys, open forests on mountain slopes;(200–)600–1700 m. Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Sichuan, Taiwan, SE Xizang, Zhejiang [India, Japan].
One of us (Jeffrey) treats this species and its varieties as synonyms
of Trichosanthes pilosa.

1a. Fruit ovoid or ovoid-oblong.
2a. Seeds oblong, middle ridge as
broad as 2 lateral locules ....... 32a. var. cucumeroides
2b. Seeds triangular, 2 lateral
locules narrow ........................... 32b. var. hainanensis
1b. Fruit globose or narrowly oblong.
3a. Fruit narrowly oblong ................ 32c. var. stenocarpa
3b. Fruit globose ........................ 32d. var. dicaelosperma
32a. Trichosanthes cucumeroides var. cucumeroides

Trichosanthes matsudae Hayata.
Fruit narrowly oblong, 7–9 × 2.5–4 cm.
Taiwan [Japan].

32d. Trichosanthes cucumeroides var. dicaelosperma (C. B.
Clarke) S. K. Chen, Bull. Bot. Res., Harbin 5(2): 118. 1985[“dicoelosperma”].
波叶栝楼 bo ye gua lou
Trichosanthes dicaelosperma C. B. Clarke in J. D. Hooker,
Fl. Brit. India 2: 609. 1879; T. ascendens C. Y. Cheng & C. H.
Yueh.
Fruit globose or oblong. Seeds compressed; 2 lateral locules suboblong.
Dense forests; 600–1200 m. Guangxi, SE Xizang [India].

33. Trichosanthes tetragonosperma C. Y. Cheng & C. H.
Yueh, Acta Phytotax. Sin. 12: 425. 1974.
方籽栝楼 fang zi gua lou
Stems slender, grooved, puberulent. Petiole 2–4 cm,
twisted, striate, puberulent; leaf blade adaxially green, narrowly
ovate, 8.5–11.5 × 4–5 cm, papery, abaxially glabrous or puberulent on veins, adaxially glabrous, apex shortly acuminate. Flowers unknown. Fruiting peduncle slender, 3–4 cm, striate, glabrous; fruit orange-yellow or red, ovoid, 6.5–11 × 4–5.5 cm,
smooth, glabrous, base rounded or obtuse, apex acute. Seeds
deep brown, 4-angled, ca. 1 × 1–1.2 cm, with narrow middle
ridge, base concave, apex triangular, 3-loculed; 2 lateral locules
broad, turgid and empty. Fr. Nov–Jan of next year.
● Forests; 1300–1600 m. SE Yunnan.

CUCURBITACEAE

46

22. CYCLANTHERA Schrader, Index Sem. Hort. Acad. Goett. 1831: 2. 1831.
小雀瓜属 xiao que gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, climbing, annual or perennial. Tendrils simple or 2- to many fid. Leaves undivided, divided, or pedately 5–7-foliolate.
Plants monoecious. Male flowers in raceme or panicle; calyx tube cup-shaped or cupular; segments 5, subulate, sometimes almost
obsolete; corolla rotate, 5-partite; segments broadly ovate-oblong, usually acute; stamen 1; filament very short; anther horizontal, 1celled, circular. Female flowers solitary, binary, or 3-fasciculate; calyx and corolla as in male flowers; ovary ovoid, 1-locular; style
very short; stigma large, orbicular. Fruit ovoid or reniform, ± fleshy, setose or setiform, rarely glabrous. Seeds few to numerous,
compressed, angled, 2-lobed at apex and base.
About 20 species: tropical America; one species (introduced) in China.

1. Cyclanthera pedata (Linnaeus) Schrader, Index Sem. Hort.
Acad. Goett. 1831: 2. 1831.
小雀瓜 xiao que gua
Momordica pedata Linnaeus, Sp. Pl. 2: 1009. 1753.
Plants annual. Stem robust, multibranched, glabrous. Petiole 5–15 cm; leaf blade 5-pedatisect, leaflets elliptic or ellipticlanceolate, median leaflet 7–16 × 2–6 cm, base cuneate, margindentate, apex acuminate; lateral leaflets 5–14 × 2–4 cm. Tendrils 2-fid. Male flowers in a panicle; peduncle 10–23 cm, 20–
50-flowered; pedicels filiform, 4–10 mm; calyx tube cupular,

ca. 2 × 3–4 mm; segments linear, ca. 2 mm; corolla yellow; segments ovate-triangular, 1.5–2 × 2–2.5 mm, puberulent; filaments connate; anthers circular, 1.5–2 mm. Female flowers:
calyx and corolla as in male flowers; ovary 2–3 mm; style 0.5–
0.7 mm. Fruit narrowly oblong to narrowly elliptic, 5–7 × 2.5–3
cm, setose, apex acuminate, 8–10-seeded. Seeds black, suboblong, 10–12 × 7–8 mm, margin dentate. Fl. and fr. May–Oct.
Cultivated. Xizang, Yunnan [cultivated in tropics of New and Old
Worlds, not known wild].
The fruit is used as a vegetable.

23. SECHIUM P. Browne, Civ. Nat. Hist. Jamaica, 355. 1756, nom. cons.
佛手瓜属 fo shou gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Chayota Jacquin; Chocho Adanson.
Herbs, scandent, perennial. Roots tuberous. Tendrils 3–5-fid. Leaf blade cordate, membranous. Plants monoecious; flowers
white. Male flowers in a raceme; calyx tube semiglobose; segments 5; corolla rotate, 5-partite; segments ovate-lanceolate, apex
acute; stamens 3, inserted on base of calyx tube; filaments connate; anthers free, one 1-celled, two 2-celled; anther cells flexuous;
pistillode absent. Female flowers solitary or binary, usually in same axil as male flowers; calyx and corolla as in male flowers;
ovary fusiform, 1-locular, setose; style short; stigma 5-lobed; lobes reflexed, with only 1 pendulous ovule. Fruit obovoid, fleshy,
sulcate at apex. Seed 1, large, ovate, compressed, woody, cotyledons large.
About five species: Mexico and Central America; one species (introduced) in China.

1. Sechium edule (Jacquin) Swartz, Fl. Ind. Occid. 2: 1150.
1800.
佛手瓜 fo shou gua
Sicyos edulis Jacquin, Enum. Syst. Pl. 32. 1760.
Stem angular-sulcate. Petiole slender, 5–15 cm, glabrous;
leaf blade suborbicular, membranous, abaxially puberulent,
adaxially slightly scabrous, base cordate, margin dentate, apex
acuminate. Male flowers 10–30 on 8–30 cm peduncle, pedicels
1–6 mm; calyx tube short; segments explanate, 5–7 mm; corolla

rotate, 12–17 mm; segments ovate-lanceolate. Female pedicel
1–1.5 cm; calyx and corolla as in male flowers; ovary obovoid,
5-angled; style 2–3 mm; stigma ca. 2 mm wide. Fruit palegreen, obovoid, 8–15 × 6–8 cm, 5-sulcate in upper part, slightly
hispid. Seed ca. 10 × 7 cm, compressed. Fl. Jul–Sep, fr. Aug–
Oct.
Commonly cultivated in S China [native to Mexico; commonly
cultivated in warm parts of the world].
The fruit is used as a vegetable.

24. MELOTHRIA Linnaeus, Sp. Pl. 1: 35. 1753.
美洲马

儿属 mei zhou ma jiao er shu

Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Landersia Macfadyen.
Herbs, climbing or trailing, annual. Leaves petiolate, usually palmately lobed. Tendrils simple. Plants monoecious. Male flowers
shortly racemose or subcorymbose; calyx campanulate to cylindrical, 5-lobed; corolla yellow, rotate; segments 5; stamens 3, inserted

CUCURBITACEAE

47

above middle of tube; anthers two 2-celled, one 1-celled; anther cells straight or slightly curved; connective not produced; rudimentary ovary subglobose. Female flowers solitary, usually coaxillary with male flowers; calyx and corolla as in male flowers; ovary
globose to fusiform, 3-locular; ovules numerous, horizontal; style surrounded at base by an annular disk; stigmas 3. Fruit globose to
oblong. Seeds numerous, compressed, smooth.
About 12 species: Neotropics; one species (introduced) in China.

1. Melothria pendula Linnaeus, Sp. Pl. 1: 35. 1753.
美洲马儿 mei zhou ma jiao er

Stems slender, setulose to glabrous. Petiole slender, retrorsely hispid, 0.9–5 cm; leaf blade adaxially dark green, abaxially paler, ovate to pentagonal, membranous to herbaceous,abaxially shortly hispid or scabrid, adaxially hispid or scabridpunctate, cordate, unlobed or shallowly to deeply 3–5-lobed,
apex obtuse to acuminate, apiculate. Male flowers 3–12 inaxillary fascicles, often with an accompanying solitary flower;
peduncle slender, 0.5–3.5 cm; pedicels slender, 1.5–9 mm; ca-

lyx campanulate, 1.5–3.5 mm, shortly pubescent to glabrous;
corolla yellow; segments spreading, obovate-oblong, 1.5–2.5mm, emarginate, apiculate; stamens 3, two 2-celled, one 1celled; filaments short, ca. 0.5 mm; anthers ellipsoid. Female
flowers solitary, often coaxillary with male flowers; pedicel
slender, 0.7–3.5 cm; calyx and corolla as in male flowers. Fruitblack when ripe, subglobose to ellipsoid, 0.8–1.9 × 0.8–1.2 cm.
Seeds pallid, ovate, compressed, minutely fimbrillate, 4–5 × 2–
2.8 mm, 0.6–0.9 mm thick. Fl. and fr. Jan–Dec.
Ruderal weeds; low elevations. Taiwan [native to the Americas (SUnited States to Argentina); introduced in SE Asia].

25. ZEHNERIA Endlicher, Prodr. Fl. Norfolk. 69. 1833.
马儿属 ma jiao er shu

Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Neoachmandra W. J. de Wilde & Duyfjes; Pilogyne Ecklon ex Schrader.
Herbs, climbing or trailing, annual or perennial. Leaves petiolate, unlobed or 3–5-lobed or -partite. Tendrils slender, simple.
Plants monoecious or dioecious. Male flowers axillary, solitary or few, long pedicellate, or in a raceme-cyme or subumbel; calyx
campanulate, 5-lobed; corolla campanulate, white or yellow-white, often becoming cream colored with age; segments 5; stamens 3,
inserted at bottom or on upper part of tube; filaments shorter to longer than anthers; anthers all 2-celled or sometimes two 2-celled
and one 1-celled, oblong or ovoid-oblong; anther cells straight or slightly curved; connective produced or not produced; rudimentary
ovary variable in form. Female flowers solitary or paired or a few in a cyme, often coaxillary with male flowers; calyx and corolla as
in male flowers; ovary globose to ellipsoid-fusiform, 3-locular; ovules few to numerous, horizontal; style surrounded at base by anannular disk; stigmas 3. Fruit globose or ellipsoid to fusiform, indehiscent. Seeds few to numerous, ovate, compressed, notsculptured.
About 55 species: Old World tropics; four species in China.

1a. Stamens inserted at bottom of tube; filaments longer than anthers.
2a. Plants monoecious; fruit globose or ovoid, red ...................................................................................................... 1. Z. bodinieri
2b. Plants dioecious; fruit oblong, purplish black ...................................................................................................... 2. Z. guamensis
1b. Stamens inserted on upper part of tube; filaments not longer than anthers.
3a. Fruit globose or oblong, less than 1.8 cm ................................................................................................................ 3. Z. japonica
3b. Fruit fusiform, attenuate, 3–4 cm, 1–1.5 cm in diam. ............................................................................................. 4. Z. wallichii
1. Zehneria bodinieri (H. Léveillé) W. J. de Wilde & Duyfjes,
Thai Forest Bull., Bot. 32: 17. 2004.
钮子瓜 niu zi gua
Melothria bodinieri H. Léveillé, Fl. Kouy-Tchéou, 122.
1914–1915; M. perpusilla Cogniaux var. subtruncata Cogniaux; Pilogyne bodinieri (H. Léveillé) W. J. de Wilde &Duyfjes.
Plants scandent. Stem and branches slender, sulcate-angular, glabrous or sparsely pubescent. Petiole slender, 2–5 cm,
glabrous; leaf blade broadly ovate or rarely triangular-ovate, 4–
12 × 3–10 cm, membranous, abaxially subglabrous, adaxiallyscabrous, basal sinus semicircular, rarely subtruncate, margin
undulate-denticulate, undivided or sometimes shortly 3–5-lobed,
apex acute or shortly acuminate. Tendrils filiform, simple, gla-

brous. Plants monoecious. Male flowers often 3–9 in a subcapitate or cymose inflorescence; peduncle slender, 1–4 cm,
glabrous; pedicels 1–4 mm; calyx tube campanulate, 2–3 × 1–2
mm, glabrous or puberulent; segments narrowly triangular, ca.
0.5 mm; corolla white; segments ovate or ovate-oblong, 2–2.5mm, abaxially puberulent, apex subacute; stamens two 2-celled,
one 1-celled, sometimes all 2-celled, inserted at base of calyx
tube; filaments ca. 2 mm, puberulent; anthers ovoid, 0.6–0.7
mm. Female flowers solitary, rarely a few in an umbel; ovary
ovoid. Fruiting pedicel slender, 0.5–1 cm, glabrous; fruit red
when ripe, baccate, globose, 1–1.4 cm in diam., smooth. Seeds
ovate-oblong, compressed, smooth, ca. 5 × 3–3.5 mm, marginate. Fl. Apr–Aug, fr. Aug–Nov.
Forest margins or roadsides on mountain slopes; 500–1000 m.
Fujian, Guangdong, Guangxi, Guizhou, Hainan, Jiangxi, Sichuan, Yunnan [India, Indonesia, Laos, Myanmar, Sri Lanka, Thailand, Vietnam].

CUCURBITACEAE

48

The Chinese record of Zehneria maysorensis (Wight & Arnott)
Arnott (FRPS 73(1): 173. 1986) is based on a misidentification of Z.
bodinieri.

2. Zehneria guamensis (Merrill) Fosberg, Smithsonian Contr.
Bot. 45: 15. 1980.
台湾马

儿 tai wan ma jiao er

Melothria guamensis Merrill, Philipp. J. Sci., C, 9: 151.
1914; M. kelungensis (Hayata) Hayata ex Makino & Nemoto;M. liukiuensis Nakai; Zehneria kelungensis Hayata; Z. liukiuensis (Nakai) E. H. Walker.
Plants scandent. Stems and branches slender, elongate, sulcate-angular, glabrous or rarely puberulent. Tendrils filiform,
simple. Petiole slender, 1.5–5 cm; leaf blade abaxially palegreen, adaxially deep green, broadly ovate, 4–8 × 4–8 cm,
membranous, scabrous, basal sinus semicircular, rarely hastate,undivided or somewhat 3–5-lobed, margin irregularly denticulate or crenate-dentate, apex acute and shortly mucronate or
shortly acuminate. Plants dioecious. Male peduncle slender,
1.5–5 cm, apex 10–30-flowered; pedicels spreading, 3–6 mm;
calyx tube broadly campanulate, ca. 3 mm, base acute; segments subulate, ca. 0.5 mm; corolla pale yellow, puberulent;segments ovate, ca. 2.5 mm, 3-veined, acute; filaments short,ca. 1 mm, puberulent; anthers ovoid, 0.8–1 mm. Female flowers
solitary or sometimes in few-flowered fascicle; pedicels 1–1.5
cm; ovary oblong. Fruit purplish black when ripe, baccate,ovate-oblong, 1–1.5(–3) cm × 7–10 mm, smooth, glabrous,
both ends obtuse. Seeds gray-white, ovate-oblong, 2–3 × ca. 2
mm, base slightly attenuate, marginate. Fl. and fr. Mar–Dec.
Forests; 400–1400 m. Guangdong, Taiwan, Yunnan [Japan; Pacific islands (Guam)].
De Wilde and Duyfjes (Blumea 51: 75. 2006) recorded Zehneria
repanda (Blume) C. Simmons (Pilogyne repanda (Blume) W. J. de
Wilde & Duyfjes) from Taiwan, on the basis of a single collection (vanSteenis 20752), which we have not seen. It differs from Z. guamensis in
its smaller, 0.5–0.8(–1) cm in diam., globose fruit. It is said to be amontane species of forest margins and scrub, at 1400–2700 m.

3. Zehneria japonica (Thunberg) H. Y. Liu, Bull. Natl. Mus.
Nat. Sci., Taichung 1: 40. 1989.
马儿 ma jiao er

Bryonia japonica Thunberg in Murray, Syst. Veg., ed. 14,870. 1784; Melothria argyi H. Léveillé; M. formosana Hayata;
M. indica Loureiro; M. japonica (Thunberg) Maximowicz ex
Cogniaux; M. leucocarpa (Blume) Cogniaux var. rubella
Gagnepain; Neoachmandra indica (Loureiro) W. J. de Wilde &Duyfjes; N. japonica (Thunberg) W. J. de Wilde & Duyfjes;
Zehneria indica (Loureiro) Keraudren; Z. omeiensis Z. Y. Zhu.
Plants scandent. Stems and branches filiform, glabrous.
Petiole slender, 2.5–3.5 cm, villous at first, glabrescent; leaf
blade abaxially pale green, adaxially dark green, polymorphic,triangular-ovate, ovate-cordate, or hastate, 3–5 × 2–4 cm, membranous, unlobed or 3–5-lobed, abaxially glabrous, adaxiallyscabrous, apex acute or rarely shortly acuminate. Plants monoecious. Male flowers solitary, or 2 or 3 in a raceme; peduncle
slender, very short, glabrous; pedicels filiform, 3–5 mm, glabrous; calyx broadly campanulate, ca. 1.5 mm, base acute or
obtuse; corolla pale yellow, puberulent; segments oblong orovate-oblong, 2–2.5 × 1–1.5 mm; stamens 3, two 2-celled andone 1-celled, sometimes all 2-celled; filaments short, ca. 0.5mm; anthers ovoid-oblong or oblong, ca. 1 mm, pubescent; anther cells curved, puberulent; connective broad, apex produced.
Female flowers solitary or rarely paired in same axil as male
flowers; pedicel filiform, 1–2 cm, glabrous; corolla broadly
campanulate, ca. 2.5 mm in diam.; segments lanceolate, 2.5–5 ×
1–1.5 mm, apex slightly obtuse; ovary narrowly ovoid, 3.5–4 ×
1–2 mm; style short, ca. 1.5 mm; stigmas 3; staminodes glandular. Fruit orange or red, oblong or attenuately ovoid, 1–1.5
cm × 5–8(–10) mm, both ends obtuse. Seeds gray-white, ovate,
3–5 × 3–4 mm, base attenuate, margin obscure. Fl. Apr–Jul, fr.
Jul–Oct.
Moist places in forests, roadsides, thickets; 500–1600 m. Anhui,Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi,
Sichuan, Taiwan, Yunnan, Zhejiang [India, Indonesia, Japan, Korea,
Nepal, Philippines, Vietnam].

4. Zehneria wallichii (C. B. Clarke) C. Jeffrey, Kew Bull. 34:
802. 1980.
锤果马儿 chui guo ma jiao er

Melothria wallichii C. B. Clarke in J. D. Hooker, Fl. Brit.
India 2: 626. 1879; Neoachmandra wallichii (C. B. Clarke) W.
J. de Wilde & Duyfjes.
Plants slender. Stems and branches glabrous. Tendrils filiform, glabrous, simple. Petiole slender, 3–4 cm, sparsely whitehirsute; leaf blade ovate or triangular, 6–7 × 6–7 cm, membranous, abaxially more scabrous, white punctate, adaxiallysparsely hirsute on veins, base truncate or with ca. 5 mm deep
sinus, margin sparsely dentate, apex acute. Plants dioecious.
Male flowers usually 7–10 in a raceme, or solitary; rachis filiform, 1–2 cm, puberulent; pedicels slender, 0.5–1 cm; calyx
tube campanulate, 1–2 × ca. 2 mm, glabrous; segments subulate, ca. 0.5 mm; corolla yellow-white; segments explanate,ovate or oblong, apex acute or slightly obtuse; stamens 3, two
2-celled, one 1-celled, inserted on base of calyx tube; filaments
short; anthers oblong, 1–1.2 mm, glabrous; anther cells erect;
connective narrow, not produced. Female flowers solitary; pedicels slender, 1–1.5 cm, puberulent; ovary fusiform, ca. 1 × 0.2
cm, subglabrous, both ends attenuate. Fruiting pedicel 1–2 cm,
sparsely hirsute; fruit fusiform, 3–4 × 1–1.5 cm, both ends
attenuate. Seeds gray-white, oblong or ovate, compressed, 5–7
× 3–4 mm, smooth, marginate. Fl. Jun–Aug, fr. Aug–Nov.
Mountain slopes; 800–1000 m. Yunnan [India, Myanmar, Thailand].

26. CUCUMIS Linnaeus, Sp. Pl. 2: 1011. 1753.
黄瓜属 huang gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, scandent or creeping, annual. Stem and branches scabrous, hispid. Tendrils slender, simple. Leaf blade almost orbicular,

CUCURBITACEAE

49

reniform, or cordate-ovate, undivided or palmately 3–7-lobed. Plants monoecious, rarely dioecious, or flowers hermaphroditic. Male
flowers fascicled or solitary; calyx tube campanulate, 5-lobed; segments subulate; corolla rotate or subcampanulate, 5-lobed; segments oblong or ovate; stamens 3, free, inserted on corolla tube; anthers oblong, one 1-celled, two 2-celled; anther cells linear,reflexed or curved; connective produced; rudimentary ovary glandular. Female flowers solitary or fascicled; calyx and corolla as inmale flowers; staminodes 3, subulate; ovary cylindric; ovules numerous, horizontal; style short; stigmas 3–5. Fruit polymorphic,
fleshy, indehiscent, smooth or verrucose. Seeds numerous, compressed, emarginate.
About 32 species: tropical and temperate regions, most species in Africa; four species in China.

1a. Fruit smooth; ovary hairy ..................................................................................................................................................... 1. C. melo
1b. Fruit and ovary muriculate or verrucose or sparsely tuberculate, not hairy.
2a. Fruit oblong or cylindric, (5–)10–50 cm ................................................................................................................... 2. C. sativus
2b. Fruit oblong, smaller than 5 cm.
3a. Fruit oblong, at least twice as long as broad, muricate ........................................................................................ 3. C. hystrix
3b. Fruit obovoid, only slightly longer than broad, sparsely obscurely tuberculate ................................................. 4. C. debilis
1. Cucumis melo Linnaeus, Sp. Pl. 2: 1011. 1753.
甜瓜 tian gua
Plants creeping. Stems and branches scabrous, puberulent,or hispid. Tendrils filiform, puberulent. Petiole 7–12 cm, setose;
leaf blade suborbicular or reniform, 6–15 × 6–15 cm, papery,
abaxially hispid, adaxially scabrous, undivided or 3–7-lobed,
dentate; sinus truncate or rounded. Plants monoecious or flowers bisexual. Male flowers 1 or 2 to several, fasciculate; pedicels filiform, 0.5–4 cm, pubescent; calyx tube narrowly campanulate, 3–8 mm, densely hispidulous to white villous; segments subulate or linear, 2–4 × 0.4–1.2 mm; corolla yellow;
tube 1–2 mm; segments ovate-oblong to broadly ovate or obovate, 3–24 × 2.5–20 mm, obtuse; anther cells flexuous. Female
flowers solitary; ovary ellipsoid or fusiform, 4–11 × 2–5 mm,
densely white lanate, villous or puberulent; style 1–2 mm; stigmas 3, 2–2.5 mm. Fruit very variable in size, shape, color, odor,
and taste. Seeds numerous, yellow-white, ovate-oblong, 4–8 ×
2.4–4 mm, smooth, base rounded, apex acute, emarginate. Fl.
and fr. May–Sep.
Farmlands, roadsides. Anhui, Jiangsu, Shandong, Xinjiang, C andE China; also cultivated throughout China [native to Old World tropics
and subtropics; introduced in New World tropics; widely cultivated intropical and temperate countries].

1a. Ovary densely villous or white lanate,with spreading hairs; (cultivated)fruit sweet ................................................... 1a. subsp. melo
1b. Ovary very shortly and densely
puberulent, hairs appressed;(cultivated) fruit not sweet .................... 1b. subsp. agrestis
1a. Cucumis melo subsp. melo甜瓜(原亚种) tian gua (yuan ya zhong)
Cucumis bisexualis A. M. Lu & G. C. Wang.
Ovary and young fruit pilose or lanate, with spreadinghairs; fruit in cultivated forms fragrant and sweet.

1b. Cucumis melo subsp. agrestis (Naudin) Pangalo in Zhukovskii, Zemledel’ch. Turtsiya, 534. 1933.
菜瓜 cai gua
Cucumis melo var. agrestis Naudin, Ann. Sci. Nat., Bot.,sér. 4, 11: 73. 1859; Bryonia callosa Rottler; C. acidus Jacquin;
C. callosus (Rottler) Cogniaux & Harms.
Ovary and young fruit sericeous with short appressedhairs; fruit in cultivated forms mostly not fragrant and not
sweet.
Farmlands, roadsides. C and E China; commonly cultivated in
China [throughout Old World tropics; introduced in New World tropics;
also commonly cultivated in E and SE Asia, rarely cultivated elsewhere].
The fruit is used as a vegetable (菜瓜 cai gua).
The forms cultivated in China belong to the ‘Conomon’ cultivar
group (Cucumis conomon Thunberg, Nova Acta Regiae Soc. Sci. Upsal.
3: 208. 1780; C. melo var. conomon (Thunberg) Makino, Bot. Mag.
(Tokyo) 16: 16. 1902).

2. Cucumis sativus Linnaeus, Sp. Pl. 2: 1012. 1753.
黄瓜 huang gua
Plants scandent or creeping. Stem and branches angular,
white hispid. Tendrils slender, simple. Petiole 10–16(–20) cm;
leaf blade broadly ovate-cordate, 7–20 × 7–20 cm, membranous, margin 3–5-angular or -lobed; lobes triangular, dentate,apex acute or acuminate; sinus half-orbicular. Plants monoecious. Male flowers fasciculate; pedicel filiform, 5–15 mm, puberulent; calyx tube narrowly campanulate, 8–10 mm, densely
white pubescent; segments subulate; corolla yellow-white, ca. 2
cm; segments oblong-lanceolate, acute; anthers 3–4 mm; connective ca. 1 mm. Female flowers solitary or fascicled; pedicels
pubescent, 1–2 cm; ovary fusiform, muricate. Fruit yellowgreen, oblong or cylindric, (5–)10–30(–50) cm, muricate, usually verrucose. Seeds white, small, narrowly ovate, 5–10 mm,
emarginate, both ends acute. Fl. and fr. summer.

Farmlands, roadsides. Anhui, Jiangsu, Shandong, Xinjiang; also
cultivated throughout China [native to Old World tropics and subtropics; introduced in New World tropics; widely cultivated in tropicaland temperate countries].

Forests, thickets, mountain slopes; 700–2000 m. Guangxi, Guizhou, Yunnan; also commonly cultivated in China [NE India, Myanmar,
Nepal, Thailand; cultivated in all tropical and temperate countries].

The fruit is used as a refreshing dessert in summer; the plants areused medicinally.

1a. Fruit oblong or cylindric, 10–30(–50) cm,
muricate, usually verrucose ......................... 2a. var. sativus

CUCURBITACEAE

50

1b. Fruit oblong or subglobose, 5–6 cm,
smooth ................................................... 2b. var. hardwickii

Commonly cultivated in China [cultivated in all tropical and temperate countries].

gin dentate, apex acute. Plants monoecious. Male flowers solitary or subfasciculate; calyx tube narrowly campanulate, ca. 6mm; segments linear, 1–2 mm; corolla yellow; segments ovateoblong, 5–6 × 3–4 mm; filaments ca. 1 mm; anthers ca. 2 mm.
Female flowers solitary; pedicels ca. 5 mm; calyx and corolla
as in male flowers; corolla 8–10 mm; ovary oblong-ovoid, ca.
10 × 4 mm, densely yellow-brown hispid; style 2–3 mm; stigmas 3-lobed. Fruit oblong, 4–5 × 1.5–2.3 cm, muricate, verrucose. Seeds narrowly ovate, 3–4 mm, both surfaces smooth,emarginate. Fl. Jun–Aug, fr. Aug–Sep.

2b. Cucumis sativus var. hardwickii (Royle) Gabaev, Ogurtsy,
47. 1932.

Forests, thickets; 800–1500 m. W Yunnan [NE India, Myanmar,
Thailand].

西南野黄瓜 xi nan ye huang gua

4. Cucumis debilis W. J. de Wilde & Duyfjes, Adansonia, sér.
3, 29: 241. 2007.

2a. Cucumis sativus var. sativus
黄瓜(原变种) huang gua (yuan bian zhong)Fruit oblong or cylindric, 10–30(–50) cm, muricate, usually verrucose.

Cucumis hardwickii Royle, Ill. Bot. Himal. Mts. 1: 220.
1835; C. sativus f. hardwickii (Royle) W. J. de Wilde &Duyfjes; C. sativus [unranked] hardwickii (Royle) Alefeld.
Fruit oblong or subglobose, 5–6 cm, smooth.
Forests, thickets, mountain slopes; 700–2000 m. Guangxi, Guizhou, Yunnan [NE India, Myanmar, Nepal, Thailand].

3. Cucumis hystrix Chakravarty, J. Bombay Nat. Hist. Soc. 50:
896. 1952.
野黄瓜 ye huang gua
Cucumis muriculatus Chakravarty.
Plants scandent. Stem and branches angular-sulcate, scabrous, white hirsute. Tendrils filiform, simple. Petiole 6–10 cm;
leaf blade broadly ovate or triangular-ovate, 6–13 × 6–10(–12)
cm, irregularly 3–5-lobed; lobes triangular, base cordate, mar-滇黄瓜 dian huang gua
Plants slender, probably annual, scandent. Stems andbranches sparsely to densely minutely aculeate-hispid. Tendrils
simple, sparsely aculeate. Petiole 3–5 cm; leaf blade subcircularin outline, 4–7 × 4–7 cm, 3–5-angular or -lobed, margin
coarsely serrate-dentate; lobes broadly triangular, base deeply
cordate, apex acute. Plants monoecious. Male flowers solitaryor paired; calyx tube turbinate-campanulate, 5.5–6 mm; segments linear, ca. 3 mm; corolla ?yellow; segments elliptic 4–6 ×
ca. 3 mm; filaments slender, ca. 0.5 mm; anthers 2.5–3 mm.
Female flowers solitary, sometimes coaxillary with male flowers; pedicels 30–35 mm; calyx and corolla probably similar tomale flowers, details unknown. Fruit green, obovoid, sparsely
obscurely tuberculate. Seeds unknown. Fl. and fr. Sep.
Yunnan [N Vietnam (ca. 1400 m)].

27. MUKIA Arnott, Madras J. Lit. Sci. 12: 50. 1840.
帽儿瓜属 mao er gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, scandent, annual, slender, setose. Petiole very short, subglabrous; leaf blade usually 3–7-lobed, base cordate. Tendrils
simple. Plants monoecious; flowers small, male flowers fascicled; female flowers usually solitary or in a few-flowered fascicle insame axil as male flowers. Male flowers: calyx campanulate; segments 5, subulate; corolla rotate; segments 5; stamens 3, free,
inserted on calyx tube; filaments short; anthers oblong, two 2-celled, one 1-celled; anther cells erect; connective slightly produced;
rudimentary ovary glandular. Female flowers: calyx and corolla as in male flowers; staminodes absent or very small; ovary ovoid,
hispid; style terete; stigma 2- or 3-lobed, ovules few, horizontal. Berry oblong or globose, small, indehiscent. Seeds horizontal, ovate,
compressed, both surfaces scabrous or smooth, marginate.
Three species: tropical and subtropical Africa, Asia, Australia; two species in China.
Renner et al. (BMC Evol. Biol. 7: 58. 2007; doi:10.1186/1471-2148-7-58) presented molecular evidence to show that Mukia is an ingroup of
Cucumis.

1a. Fruit globose, fruit coat thickening; seed surfaces scrobiculate, indistinctly marginate ................................... 1. M. maderaspatana
1b. Fruit oblong, fruit coat not thickening; seed surfaces indistinctly scrobiculate, marginate ......................................... 2. M. javanica
1. Mukia maderaspatana (Linnaeus) M. Roemer, Fam. Nat.
Syn. Monogr. 2: 47. 1846.
帽儿瓜 mao er gua

rella Linnaeus f.; Coccinia cordifolia (Linnaeus) Cogniaux;
Melothria althaeoides (Seringe) Nakai; M. maderaspatana(Linnaeus) Cogniaux; Mukia althaeoides (Seringe) M. Roemer;
M. scabrella (Linnaeus f.) Arnott.

Cucumis maderaspatanus Linnaeus, Sp. Pl. 2: 1012. 1753;
Bryonia althaeoides Seringe; B. cordifolia Linnaeus; B. scab-

Plants scandent or prostrate, all parts densely yellowbrown hispid. Stems many branched, robust, sulcate. Leaf blade

CUCURBITACEAE

somewhat rigid, ovate or ovate-cordate, 5–9 × 5–9 cm, usually
3–5-lobed, median lobe ovate-triangular, both surfaces scabrous, base cordate, margin irregularly denticulate, apex slightly
obtuse. Tendrils moderately robust. Male flowers fascicled;
pedicels short or absent; calyx tube campanulate, ca. 2 × 1.5mm; segments reflexed, subulate, 1–1.5 × ca. 1 mm; corolla
yellow; segments ovate-oblong, ca. 2 × 0.5 mm, apex obtuse;
filaments short, ca. 0.5 mm, slightly pilose; anthers oblong, ca.
1 mm, ciliate; connective distinct; rudimentary ovary globose.
Female flowers solitary or 3–5-fasciculate. Fruiting pedicels
extremely short; fruit dark red, globose, ca. 1 cm in diam.,smooth. Seeds ovate, ca. 4 × 2.5 mm, ca. 2.5 mm thick, both
surfaces scrobiculate, indistinctly marginate. Fl. Apr–Aug, fr.
Aug–Dec.
Rocky mountain slopes, thickets; 400–1700 m. Guangdong,
Guangxi, Guizhou, Taiwan, Yunnan [tropical Africa and Asia, Australia].

2. Mukia javanica (Miquel) C. Jeffrey, Hooker’s Icon. Pl. 37:
t. 3661. 1969.
爪哇帽儿瓜 zhua wa mao er gua
Karivia javanica Miquel, Fl. Ned. Ind. 1(1): 661. 1856;
Cucumis javanicus (Miquel) Ghebretinsae & Thulin; Melothria

51

assamica Chakravarty; M. assamica var. scabra Chakravarty;
M. javanica (Miquel) Cogniaux.
Plants scandent. Stem and branches slender, hispid. Petiole
shortly hispid; leaf blade ovate-cordate, 4–6 × 3–5(–7) cm,
usually 3–5-lobed, median lobe ovate-triangular, both surfaces
shortly hispid, margin dentate, apex acuminate. Tendrils slender, hispid. Male flowers 2 to several, fasciculate, in leaf axils;
pedicel very short, 2–3 mm, shortly hispid; calyx tube cupular,
2–2.5 mm; segments erect, subulate, 1.5–2.5 mm; corolla yellow; segments ovate-oblong, 3–4 mm, apex obtuse; stamens inserted on calyx tube; filaments ca. 0.5 mm, puberulent; anthers
1.5–2 mm; connective produced. Female flowers fasciculate insame axil as male flowers; calyx tube cupular; segments triangular, ca. 0.5 × 0.3 mm; corolla segments ovate-oblong, ca. 2.5
× 1.5 mm, 3-veined; ovary ovoid, 3–4 × 1.5–2.5 mm, densely
hispid; style ca. 1 mm, apex 3-lobed; staminodes glandular.
Fruiting pedicel 2–4 mm, densely hispid; fruit red, oblong, 1–
1.5 × ca. 1 cm. Seeds ovate, ca. 4 × 2.5 mm, margin prominent. Fl. Apr–Jul, fr. Jul–Oct.
Forests, grasslands on mountain slopes; 500–1200 m. Guangdong,
Guangxi, Taiwan, Yunnan [India, Indonesia, Philippines, Thailand,
Vietnam].

28. SCOPELLARIA W. J. de Wilde & Duyfjes, Blumea 51: 297. 2006.
云南马儿属 yun nan ma jiao er shu

Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Scopella W. J. de Wilde & Duyfjes, Blumea 51: 34. 2006, not Mains (1939) [Fungi].
Herbs, climbing, annual or perennial. Leaves petiolate, unlobed or lobed. Tendrils simple, hairy. Plants monoecious. Male flowers shortly racemose; calyx campanulate, 5-lobed; corolla yellow; segments 5; stamens 3, inserted on throat of tube; filaments longer
than anthers; anthers all 2-celled, anther cells straight; connectives not produced; rudimentary ovary subglobose. Female flowers 1 or
2, often coaxillary with male flowers; calyx and corolla as in male flowers; ovary ellipsoid-globose, 3-locular; ovules numerous,
horizontal; style surrounded at base by an annular disk; stigmas 3. Fruit globose to fusiform, indehiscent. Seeds numerous, ovoidellipsoid, somewhat compressed, faces scrobiculate.
Two species: SE Asia and W Malaysia; one species in China.
One of us (Lu) does not recognize “micro-genera” such as this; however, for the purposes of this Flora, the treatment by the co-author (Jeffrey)
is followed.

1. Scopellaria marginata (Blume) W. J. de Wilde & Duyfjes,
Blumea 51: 297. 2006.
云南马儿 yun nan ma jiao er

Bryonia marginata Blume, Bijdr. 924. 1826; Melothria
marginata (Blume) Cogniaux; Scopella marginata (Blume) W.
J. de Wilde & Duyfjes; Zehneria marginata (Blume) Keraudren.
Plants scandent, annual. Stem and branches glabrous, angular-sulcate. Petiole slender, 2–3.5 cm, white hirsute; leaf blade
ovate-cordate or oblong-hastate, 6–10 × 4–10 cm, membranous,
usually 3-lobed; median lobe larger, narrowly triangular or lanceolate-triangular, 5–7 × 1.5–2.5 cm, apex acute; lateral lobes
smaller, triangular, 2.5–3.5 × 2–2.5 cm, scabrous on both
surfaces, white setose, base truncate or slightly concave, mar-gin callous-dentate, apex obtuse. Tendrils slender. Male flowers
usually 5–15 in a 4–6 cm raceme; pedicels only 2–6 mm, puberulent; calyx tube narrowly campanulate, ca. 1 mm, hirsute;segments subulate, 0.3–0.5 mm; corolla yellow, pubescent; segments ovate-triangular, ca. 1 mm; anthers attenuate, glabrous;
connective not produced; pistillodes glandular. Female flowers
solitary in axils; pedicels filiform, 3–4 cm; calyx and corolla as
in male flowers; ovary narrowly oblong, puberulent; styles
short; stigmas enlarged. Fruiting pedicel slender, 3–4 cm, glabrous; fruit ovoid or subglobose, 1.5–1.8 cm × 5–12 mm, baseslightly obtuse or attenuate. Seeds ovate, ca. 4 × 2.2 mm, ca. 1
mm thick, base obtuse-rounded, both surfaces scrobiculate,margin prominent.
Sparse forests or roadsides on mountain slopes; 600–800 m. Yunnan [Cambodia, Indonesia, Laos, Myanmar, Philippines, Thailand, Vietnam].

CUCURBITACEAE

52

29. DIPLOCYCLOS (Endlicher) T. Post & Kuntze, Lex. Gen. Phan. 178. 1903 [“Diplocyclus”].
毒瓜属 du gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Bryonia [unranked] Diplocyclos Endlicher, Prodr. Fl. Norfolk. 68. 1833.
Herbs, scandent. Tendrils 2-fid. Leaf blade palmately 5-lobed. Plants monoecious. Flowers small, fasciculate, often male andfemale flowers together in axils. Male flowers: calyx tube broadly campanulate; segments short; corolla broadly campanulate; segments ovate; stamens 3, free, inserted on calyx tube; filaments short; anthers ovoid, one 1-celled, other 2-celled; anther cells linear,slightly flexuous. Female flowers: calyx and corolla as in male flowers; staminodes 3; ovary globose or ovoid, with 3 placentas;
ovules few, horizontal; style slender; stigmas 3, 2-lobed. Fruit baccate, globose or ovoid. Seeds strongly margined, surfaces slightly
scrobiculate at middle.
Four species: tropical Africa, Asia, Australia; one species in China.

1. Diplocyclos palmatus (Linnaeus) C. Jeffrey, Kew Bull. 15:352. 1962.
毒瓜 du gua
Bryonia palmata Linnaeus, Sp. Pl. 2: 1012. 1753; B.
affinis Endlicher; Bryonopsis affinis (Endlicher) Cogniaux; B.
laciniosa (Linnaeus) Naudin var. erythrocarpa Naudin; B.
laciniosa var. walkeri Chakravarty; Ilocania pedata Merrill.
Root tuberous. Stems slender, glabrous. Tendrils glabrous.
Petiole 4–6 cm, slightly puberulent; leaf blade broadly ovate, 8–
12 × 8–12 cm, membranous, palmately 5-lobed; middle lobe
oblong-lanceolate, 8–10 × 2–3.5 cm, apex acuminate or obtuse;lateral lobes short, lanceolate or oblong-lanceolate, both surfaces subglabrous, sinus rounded. Male and female flowers

usually fasciculate in same axil. Male flowers: pedicel slender,
5–15 mm; calyx tube ca. 2 × 5–6 mm; segments subulate, 0.5–1mm; corolla green-yellow, ca. 7 mm in diam.; segments ovate,ca. 2 × 0.5–1 mm; stamens 3; filaments 1–1.5 mm; anthers
ovoid, ca. 2 mm; anther cells reflexed. Female flowers: calyx
and corolla as in male flowers; ovary ovoid, smooth. Fruit yellow-green to red, narrowly white striped, globose, indehiscent,
14–18 mm in diam., smooth. Seeds few, brown, ovate, ca. 5 × 3
mm. Fl. Mar–Aug, fr. Sep–Dec.
Forests, thickets; ca. 1000 m. Guangdong, Guangxi, Hainan,
Taiwan [Bhutan, Cambodia, India, Japan (Ryukyu Islands), Malaysia,
Nepal, Philippines, Sri Lanka, Thailand, Vietnam; tropical Africa,
Australia].
The fruit and roots contain toxin.

30. COCCINIA Wight & Arnott, Prodr. Fl. Ind. Orient. 1: 347. 1834.
红瓜属 hong gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Cephalandra Schrader.
Herbs, climbing. Roots tuberous. Branches glabrous or slightly scabrous. Tendrils simple, rarely 2-fid. Leaf blade angled or
divided. Plants dioecious or rarely monoecious. Male flowers solitary or in a cyme or raceme; calyx tube short, campanulate or turbinate; segments 5; corolla campanulate; segments 5; stamens 3, inserted at base of calyx tube; filaments connate; anthers connivent,one 1-celled, two 2-celled; anther cells reflexed; connective not produced. Female flowers solitary; calyx and corolla as in male
flowers; staminodes 3, oblong or subulate; ovary ovoid, oblong, or linear; placentas 3; ovules numerous, horizontal; style filiform;
stigma 3-lobed. Fruit ovoid or oblong, baccate, indehiscent. Seeds numerous, compressed, marginate.
About 20 species: mainly distributed in tropical Africa; one species in China.

1. Coccinia grandis (Linnaeus) Voigt, Hort. Suburb. Calcutt.
59. 1845.
红瓜 hong gua
Bryonia grandis Linnaeus, Syst. Nat., ed. 12, 2: 640;
Mant. Pl. 1: 126. 1767; Cephalandra indica Naudin, nom. illeg.
superfl.; Coccinia indica Wight & Arnott, nom. illeg. superfl.
Roots robust. Stem slender, slightly woody, manybranched, angular, glabrous. Tendrils filiform, glabrous, simple.
Petiole slender, 2–5 cm; leaf blade broadly cordate, 5–10 × 5–
10 cm, usually 5-angled or 5-lobed, base with several glands,
apex obtuse. Plants dioecious; flowers solitary. Male pedicel

slender, 2–4 cm, glabrous; calyx tube broadly campanulate, 4–5mm; segments linear-lanceolate, ca. 3 mm; corolla white orslightly yellow, 2.5–3.5 cm; segments ovate, outside glabrous,
inside pubescent; filaments and anthers connate; filaments 2–3mm; anthers subglobose, 6–7 mm. Female pedicel slender, 1–3
cm; staminodes 3, 1–3 mm, nearly subulate, villous at base;ovary fusiform, 12–15 mm; style 6–7 mm; stigmas 3, 5–6 mm.
Fruit red when mature, fusiform, ca. 5 × 2.5 cm. Seeds yellow,
oblong, 6–7 × 2.5–4 mm, ca. 1.5 mm thick, both surfaces
densely punctate, apex rounded. Fl. and fr. summer.
Thickets on mountain slopes and in forests; 100–1100 m. Guangdong, Guangxi, Yunnan [tropical Africa and Asia].

CUCURBITACEAE

53

31. LAGENARIA Seringe, Mém. Soc. Phys. Genève 3(1): 25, t. 2. 1825.
葫芦属 hu lu shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, scandent, softly pubescent. Tendrils 2-fid. Petiole apex 2-glandular; leaf blade ovate-cordate or reniform. Plants dioecious or monoecious; flowers solitary or male flowers in racemes, white. Male flowers: pedicels elongate; calyx tube narrowly campanulate or funnelform; segments 5; corolla segments 5, oblong-obovate, apex retuse; stamens 3; filaments free; anthers included,slightly coalescent or free, oblong, one 1-celled, others 2-celled; anther cells sigmoid-flexuous; connective not produced; pistillode
glandular. Female flowers: pedicels short; calyx tube cupular; calyx and corolla as in male flowers; ovary ovoid or cylindric; ovules
numerous, horizontal; placentas 3; styles short; stigmas 3, 2-lobed. Fruit polymorphic, indehiscent, fleshy, woody when mature.
Seeds numerous, obovate, compressed, distinctly marginate.
Six species: mainly distributed in tropical Africa; one species (introduced) in China.

1. Lagenaria siceraria (Molina) Standley, Publ. Field Mus.
Nat. Hist., Bot. Ser. 3: 435. 1930.
葫芦 hu lu
Cucurbita siceraria Molina, Sag. Stor. Nat. Chili, 133.
1782; Cucumis mairei H. Léveillé; Cucurbita lagenaria Linnaeus; C. leucantha Duchesne, nom. illeg. superfl.; Lagenaria
leucantha Rusby; L. leucantha var. clavata Makino; L. leucantha var. depressa (Seringe) Makino; L. leucantha var. makinoi Nakai; L. leucantha var. microcarpa (Naudin) Nakai; L.
microcarpa Naudin; L. siceraria var. depressa (Seringe) H.
Hara; L. siceraria var. microcarpa (Naudin) H. Hara; L. vulgaris Seringe; L. vulgaris subsp. asiatica Kobjakova; L. vulgaris var. depressa Seringe; L. vulgaris var. microcarpa Matsumura & Nakai.
Plants annual. Stem and branches sulcate-angular, pubes-

cent. Tendrils filiform, puberulent. Petiole 16–20 cm, with 2
glands at apex; leaf blade ovate-cordate or reniform-ovate, 10–
35 × 10–35 cm, undivided or 3–5-lobed, both surfaces puberulent, palmately 5–7-veined, base cordate, margin irregularly
dentate. Plants monoecious; flowers solitary. Male flowers:
pedicels slender; calyx tube funnelform, ca. 2 cm; segments
lanceolate, ca. 5 mm; corolla yellow; segments 3–4 × 2–3 cm;
filaments 3–4 mm; anthers oblong, 8–10 mm. Female flowers:
calyx segments and corolla as in male flowers; ovary densely
pubescent; stigma enlarged, 2-lobed. Fruit various in size andshape, woody when mature. Seeds white, obovate or triangular,
ca. 20 mm, apex truncate. Fl. summer, fr. autumn.
Commonly cultivated in China [cultivated in all warmer areas of
the world].
The fruit is used as a vegetable when young; woody when mature,
it is used as various vessels.

32. CITRULLUS Schrader ex Ecklon & Zeyher, Enum. Pl. Afric. Austral. 2: 279. 1836,
nom. cons.
西瓜属 xi gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Anguria Miller.
Herbs, climbing, annual or perennial. Stem and branches scabrous, robust. Tendrils 2- or 3-fid, rarely simple. Leaves orbicularor triangular-ovate, deeply 3–5-partite; segments lobulate or dissected. Plants monoecious; flowers solitary, rarely fasciculate. Male
flowers: calyx tube broadly campanulate, 5-lobed; corolla rotate or broadly campanulate, deeply 5-partite; segments ovate-oblong;stamens 3, inserted on base of calyx tube; filaments free, short; anthers free or slightly coherent, one 1-celled, others 2-celled; anthercells linear, sigmoid-flexuous; connective dilated, not produced; rudimentary ovary glandlike. Female flowers: calyx and corolla as
in male flowers; staminodes 3, setiform or ligulate; ovary ovoid, with 3 placentas; ovules numerous, horizontal; style short,columnar; stigmas 3, thick, reniform, nearly 3-lobed. Fruit globose or oblong, fleshy or dry, indehiscent, many seeded. Seeds oblong,compressed, smooth, marginate or emarginate.
Four species: tropical and S Africa, SW Asia, E Mediterranean region; one species (introduced) in China.

1. Citrullus lanatus (Thunberg) Matsumura & Nakai, Cat.
Sem. Spor. Hort. Bot. Univ. Imp. Tokyo 30: no. 854. 1916.
西瓜 xi guaMomordica lanata Thunberg, Prodr. Pl. Cap. 13. 1794;
Citrullus colocynthis Schrader var. lanatus (Thunberg) Matsumura & Nakai; C. edulis Spach; C. vulgaris Schrader ex Ecklon& Zeyher; Colocynthis citrullus (Linnaeus) Kuntze; Cucumis
citrullus (Linnaeus) Seringe; Cucurbita citrullus Linnaeus.

Plants annual. Stem and branches robust, sulcate-angular,
villous. Tendrils puberulent, 2-fid. Petiole 3–12 cm, densely pubescent; leaf blade white-green, triangulate-ovate, 8–20 × 5–15
cm, both surfaces hispid, 3-partite; segments lobulate, base cordate, sinus semicircular, apex acute or acuminate. Flowers solitary. Male flowers: pedicel 3–4 cm, villous; calyx tube densely
villous; segments narrowly lanceolate, 2–3 mm; corolla paleyellow, 2.5–3 cm in diam.; segments ovate-oblong, 1–1.5 ×
0.5–0.8 cm; stamens nearly free. Female flowers: calyx andCUCURBITACEAE

54

corolla as in male flowers; ovary 5–8 mm, densely villous;
stigmas 3, reniform. Fruit globose or oblong, smooth. Seeds
numerous, color various, ovate, 1–1.5 × 0.5–0.8 cm. Fl. and fr.
Apr–Oct.

Commonly cultivated in China [native to S Africa; cultivated in
all warmer areas of the world].
The fruit are important “water fruit” in summer; the seeds contain
edible oil; the fruit coat is used medicinally.

33. SOLENA Loureiro, Fl. Cochinch. 2: 477, 514. 1790.
茅瓜属 mao gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Herbs, scandent, perennial. Roots tuberous. Stem and branches slender, subglabrous. Tendrils simple, glabrous. Petiole very
short or almost obsolete; leaf blade polymorphic, very variable, base cordate or hastate. Plants dioecious or monoecious. Male
flowers in umbelliform or cymose inflorescence; calyx tube campanulate; segments 5, ± subulate; corolla yellow or yellow-white;segments triangular; stamens 3, two 2-celled, one 1-celled; filaments short; anthers oblong; anther cells conduplicate or curved.
Female flowers solitary; ovary oblong; ovules few, horizontal; staminodes 3. Fruit oblong or ovoid, indehiscent. Seeds few, globose.
Three species: S and SE Asia; one species in China.

1. Solena heterophylla Loureiro, Fl. Cochinch. 2: 514. 1790.
茅瓜 mao gua
Roots fusiform, 1.5–2 cm in diam. Stem and branches glabrous. Petiole slender, 4–10 mm, puberulent at first, glabrescent; leaf blade very variable, ovate, oblong, ovate-triangular, or
hastate, undivided or 3–5-lobed, leathery; lobes oblong-lanceolate, lanceolate, or triangular, 8–12 × 1–5 cm, abaxially densely
setose or almost glabrous, adaxially densely setose or scabrous,
base cordate, margin entire or dentate, apex obtuse or acuminate. Tendrils slender. Male flowers umbellate or subumbellate;peduncle very short, apically 10–20-flowered; pedicels 2–8mm; calyx tube 3–5 mm, ca. 3 mm in diam.; segments subulate,
0.2–0.4 mm; corolla yellow or yellow-white; segments triangular, 1–1.5 mm, apex obtuse or acute; filaments filiform, ca. 3
mm; anther cells curved or conduplicate, puberulent. Female
flowers usually solitary; pedicel 2–10 mm, puberulent; calyx
and corolla as in male flowers; ovary ovoid, 2.5–3.5 × 2–3 mm;stigmas 3. Fruit red-brown, broadly ovoid, oblong, or subglobose, 2–6 × 2–5 cm. Seeds gray-white or gray-brown, suborbicular or obovate, 5–7 × 5–6.5 mm, smooth or slightly tuberculate. Fl. May–Aug, fr. Jun–Nov.
Mixed forests, thickets, grasslands, roadsides, mountain slopes;
600–2600 m. Fujian, Guangdong, Guangxi, Guizhou, Jiangxi, Sichuan,
Taiwan, Xizang, Yunnan [NE Afghanistan, Bhutan, India, Indonesia(Java), Malaysia, Myanmar, Nepal, Pakistan, Thailand, Vietnam].
The roots are used medicinally.

1a. Plants dioecious; leaves polymorphic;
anther cells curved ......................... 1a. subsp. heterophylla

1b. Plants monoecious; leaves palmately
5-lobed, segments lanceolate; anthercells conduplicate ........................... 1b. subsp. napaulensis
1a. Solena heterophylla subsp. heterophylla
茅瓜(原亚种) mao gua (yuan ya zhong)
Bryonia hastata Loureiro; Melothria delavayi Cogniaux;
Solena delavayi (Cogniaux) C. Y. Wu.
Plants dioecious. Anther cells curved. Fruit oblong or subglobose, 2–6 × 2–5 cm. Fl. May–Aug, fr. Aug–Nov.
Mixed forests, thickets, grasslands, roadsides on mountain slopes;
600–2600 m. Fujian, Guangdong, Guangxi, Guizhou, Jiangxi, Sichuan,
Taiwan, Xizang, Yunnan [NE Afghanistan, India, Indonesia (Java),
Malaysia, Myanmar, Nepal, Thailand, Vietnam].
The roots are used medicinally.

1b. Solena heterophylla subsp. napaulensis (Seringe) W. J. de
Wilde & Duyfjes, Blumea 49: 75. 2004.
西藏茅瓜 xi zang mao gua
Bryonia napaulensis Seringe in Candolle, Prodr. 3: 307.
1828.
Plants monoecious. Anther cells conduplicate. Fruit
broadly ovoid, 2.5–3 × ca. 2.5 cm, glabrous, apex and baseobtuse. Fl. May–Jun, fr. Jun–Aug.
Mountain slopes, grasslands, thickets; 2000–2300 m. SE Xizang,
Yunnan [N India, Myanmar, Nepal].

34. BENINCASA Savi, Bibliot. Ital. Giorn. Lett. 9: 158. 1818.
冬瓜属 dong gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Camolenga Post & Kuntze.
Herbs, creeping, annual, all parts densely hispid. Leaves palmately 5-lobed; petiole eglandular. Tendrils 2- or 3-fid. Plants usually monoecious; flowers solitary. Male flowers: calyx tube broadly campanulate; segments reflexed, almost leaflike, dentate; corolla
rotate, usually 5-lobed; segments obovate, entire; stamens 3, free, inserted on corolla tube; filaments short; anthers one 1-celled, two
2-celled; anther cells very sigmoid-flexuous; connective margined; pistillode glandlike. Female flowers: calyx and corolla as in male

CUCURBITACEAE

55

flowers; staminodes 3; ovary ovoid; ovules numerous, horizontal; style inserted on disk; stigmas 3, expanded, 2-lobed. Fruit large,
oblong, baccate, terete, hispid, glaucous, indehiscent. Seeds numerous, ovoid-oblong, compressed, margin tumid.
One species: cultivated in tropical and subtropical regions of the world, including China.

1. Benincasa hispida (Thunberg) Cogniaux in A. Candolle &C. Candolle, Monogr. Phan. 3: 513. 1881.
冬瓜 dong gua
Cucurbita hispida Thunberg, Nova Acta Regiae Soc. Sci.
Upsal. 4: 38. 1783; Benincasa cerifera Savi; B. hispida var.
chieh-qua F. C. How; B. pruriens (Solander ex Seemann) W. J.
de Wilde & Duyfjes; B. pruriens f. hispida (Thunberg) W. J. de
Wilde & Duyfjes; C. pruriens Solander ex Seemann; Lagenaria
leucantha Rusby var. hispida (Thunberg) Nakai; L. siceraria(Molina) Standley var. hispida (Thunberg) H. Hara; L. vulgaris
Seringe var. hispida (Thunberg) Nakai.
Plants creeping or scandent. Stem and branches yellowbrown hispid and villous, sulcate-angular. Petiole robust, 5–
20 cm, yellow-brown hispid and villous; leaf blade reniformorbicular, 15–30 cm wide, 5–7-lobed; lobes broadly triangularor ovate, both surfaces scabrous, abaxially densely pubescent,adaxially puberulent, base deeply cordate, sinus almost rounded,margin dentate, apex acute. Male flowers: pedicel 5–15 cm,
densely hispid and villous; bract ovate or broadly oblong, 6–10
mm, apex acute; calyx tube 12–15 mm in diam., densely hispidvillous; segments 8–12 mm; corolla yellow; segments 3–6 ×
2.5–3.5 cm, both surfaces pubescent; filaments 2–3 mm, base
expanded; anthers ca. 5 mm. Female flowers: pedicel less than
5 cm, densely yellow-brown hispid and villous; ovary ovoid or
cylindric, 2–4 cm, densely hispid; style 2–3 mm; stigmas 12–15
mm. Fruit 50–60 × 10–25 cm. Seeds white or pale yellow, 10–
11 × 5–7 mm, ca. 2 mm thick, marginate. Fl. Jun–Sep, fr. Jul–
Nov.
Commonly cultivated in China [native origin uncertain; cultivatedthroughout the tropics, especially in Asia].
The fruit is eaten as a vegetable; the seeds are used medicinally.

35. CUCURBITA Linnaeus, Sp. Pl. 2: 1010. 1753.
南瓜属 nan gua shu
Lu Anmin (路安民 Lu An-ming); Charles Jeffrey
Pepo Miller.
Herbs, climbing or prostrate, annual. Stem and branches robust. Tendrils 2- to many fid. Leaf blade lobed, base cordate. Plantsmonoecious; flowers solitary, yellow. Male flowers: calyx tube campanulate, rarely elongate; segments 5, lanceolate or leaflike at
apex; corolla campanulate, 5-lobed; stamens 3; filaments free; anthers connected into a head, one 1-celled, two 2-celled; anther cells
linear, reflexed; connective not produced; pistillode absent. Female peduncle short; calyx and corolla as in male flowers; staminodes
3, broadly triangular; ovary oblong or globose, with 3–5-placentas; ovules numerous, horizontal; style short; stigmas 3–5-lobed orbifurcate. Fruit large, fleshy, indehiscent. Seeds numerous, compressed, smooth.
About 15 species: warm regions of the Americas, also cultivated in tropical, subtropical, and temperate regions; three species (all introduced) in
China.

1a. Calyx segments linear, apex leaflike; fruiting pedicel conspicuously enlarged at apex; seed margin thickened ....... 3. C. moschata
1b. Calyx segments linear or linear-lanceolate, apex not leaflike; fruiting pedicel not strongly enlarged at apex.
2a. Leaf blade triangular or ovate-triangular, irregularly 5–7-lobed; calyx segments linear-lanceolate; fruiting
pedicel angular-sulcate, slightly thickened at apex; seed marginate and obtuse .......................................................... 1. C. pepo
2b. Leaf blade reniform or orbicular, almost entire or dentate; calyx segments lanceolate; fruiting pedicel not
angular-sulcate, not thickened at apex; seed margin obtuse or ± marginate ........................................................... 2. C. maxima
1. Cucurbita pepo Linnaeus, Sp. Pl. 2: 1010. 1753.
西葫芦 xi hu lu
Plants climbing. Stem setose and scabrous-hairy. Petiole
setose, 6–9 cm; leaf blade triangular or ovate-triangular, both
surfaces scabrous-hairy, base cordate, margin irregularly dentate, apex acute. Tendrils many fid. Plants monoecious; flowers
solitary; pedicel angular, 3–6 cm, yellow-brown setose; calyx
tube 5-angled; segments linear-lanceolate; corolla campanulate,ca. 5 × 3 cm; segments erect, apex acute; stamens 3; filaments
ca. 15 mm; anthers connivent, ca. 10 mm; ovary ovoid, 1locular. Fruiting pedicel robust, conspicuously angular-sulcate,apex slightly thickened; fruit variable in shape and size. Seeds
numerous, white, ovate, ca. 20 mm, marginate and obtuse. Fl.
and fr. May–Nov.

Commonly cultivated throughout China [native to North America;
cultivated in tropical and temperate regions].
The fruit is used as a vegetable.

2. Cucurbita maxima Duchesne, Essai Hist. Nat. Courges, 7,
12. 1786.
笋瓜 sun gua
Cucurbita pepo Linnaeus var. maxima (Duchesne) Delile.
Plants climbing. Stem robust, white setose. Petiole 15–20
cm, densely setose; leaf blade reniform or orbicular-reniform,
15–25 cm, both surfaces setose, base cordate, margin almost
entire or dentate, apex obtuse. Tendrils usually many fid,slightly setose. Male pedicel 10–20 cm, puberulent; calyx tube
campanulate; segments linear-lanceolate, 18–20 mm, densely

CUCURBITACEAE

56

white setose; corolla tubular; segments reflexed, ovate-orbicular, 2–3 cm, margin rugose, apex obtuse; stamens 3; filaments
connivent, 5–7 mm, almost glabrous or puberulent at base;anthers connivent. Ovary ovoid; style short; stigmas 2-lobed.
Fruiting pedicel short, cylindric, not angular-sulcate, apex not
enlarged; fruit variable in shape, size, and color. Seeds compressed, margin obtuse or marginate. Fl. and fr. Apr–Nov.
Commonly cultivated in China [native to South America; cultivated in tropical and temperate regions].
The fruit is used as a vegetable; the seeds contain oil.

3. Cucurbita moschata Duchesne, Essai Hist. Nat. Courges, 7,
15–16. 1786.
南瓜 nan gua
Cucurbita pepo Linnaeus var. moschata (Duchesne) Duchesne.

Plants prostrate. Stem to 2–5 m, densely white setose.
Petiole robust, 8–19 cm, setose; leaf blade broadly ovate orovate-orbicular, 12–25 × 20–30 cm, 5-angled or 5-lobed; lobes
triangular, adaxially yellow-white setose and velvety. Male calyx tube campanulate, 5–6 mm; segments linear, 1–1.5 cm, pubescent, enlarged or leaflike at apex; corolla campanulate, ca. 8
× 6 cm; segments with revolute margin, rugose, apex acute;stamens 3; filaments 5–8 mm, glandular; anthers connivent, ca.
15 mm. Ovary 1-locular; style short; stigma enlarged, 2-lobed.
Fruiting pedicel robust, 5–7 cm, angular-sulcate, strongly enlarged at apex; fruit variable in shape, size, and color. Seeds numerous, gray-white, ovate or oblong, 10–15 × 7–10 mm, margin thickened. Fl. and fr. Apr–Nov.
Commonly cultivated throughout China [native to Central America; cultivated worldwide].
The fruit is used as a vegetable; the seeds are edible.